Domestication of plants in the Old World: the origin and spread of domesticated plants in Southwest Asia, Europe, and the Mediterranean Basin [4th edition] 9780199549061, 0199549060, 9780199688173, 0199688176

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Domestication of Plants in the Old World

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Domestication of Plants in the Old World The origin and spread of domesticated plants in south-west Asia, Europe, and the Mediterranean Basin Fourth Edition Daniel Zohary Professor Emeritus Department of Evolution, Systematics and Ecology The Hebrew University of Jerusalem, Israel

Maria Hopf* Formerly Head of the Botany Department, Römisch-Germanisches Zentralmuseum, Mainz, Germany and

Ehud Weiss Senior Lecturer Archaeobotanical Laboratory The Institute of Archaeology The Martin (Szusz) Department of Land of Israel Studies and Archaeology Bar-Ilan University, Ramat-Gan, Israel and Kimmel Center for Archaeological Sciences Weizmann Institute of Science, Rehovot, Israel *deceased

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Great Clarendon Street, Oxford ox2 6dp Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York © Daniel Zohary, Maria Hopf, and Ehud Weiss 2012 The moral rights of the authors have been asserted Database right Oxford University Press (maker) First edition published 1988 Second edition published 1993 Third edition published 2000 Fourth edition published 2012 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by SPI Publisher Services, Pondicherry, India Printed and bound by CPI Group (UK), Ltd, Croydon, CR0 4YY ISBN 978–0–19–954906–1 1 3 5 7 9 10 8 6 4 2

Dedicated to the memory of Dr Maria Hopf

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Preface to the fourth edition

Sadly, Dr Maria Hopf passed away since the publication of the third edition. As a token of our enormous appreciation of her warm, kind personality, her work in the compilation and updating of this book, and her other scientific achievements, we dedicate this edition to her memory. A considerable amount of new data and research avenues came to light over the past decade, triggering the preparation of this new edition. Many new archaeological sites, some from untreated regions, have been excavated and analyzed archaeobotanically, and new research tools developed. These tools, mostly in the field of molecular analysis of living plants (and animals), drove many enthusiastic research groups to explore questions relating to the domestication process and the relationship between crops and their wild ancestors. Such new representative data are included.

Main changes in the fourth edition The field of radiocarbon dating improved dramatically in the last generation, and dating labs are collaborating more extensively with each other and research. Although not all sites mentioned in this edition were dated to the same methodical degree, we now have a much more accurate baseline for comparison. Therefore, this edition makes use of calibrated dates, when they are available. As a result, our ability to compare sites has improved, and greater accuracy gained in our understanding of the movement of south-west Asian assemblage crops beyond their ‘core area’. In an attempt to improve our knowledge about the transfer of the Neolithic package to Europe, Asia, and Africa, we consulted our colleagues for the most representative sites in each country. As a

result, Chapter 10 ‘Plant remains in representative archaeological sites’, has been significantly updated. Thirty-eight sites were deleted and sixty-four were added (36% new sites out of a total of 179 sites). North Africa has been added to the geographical coverage of this edition, but the Indian subcontinent was excluded. Many research projects flourished in the Indian subcontinent and eastern Asia in the last decade, and we believe that these regions deserve specific treatment. We have made the text easier to read, with renewed artwork, figures, tables, and maps. Nineteen colour plates and eight new black and white figures have been added, as well as a new map, (Map 2) which summarizes the spread of the south-west Asian Neolithic crop assemblage in Europe, west Asia, and North Africa. The concluding chapter now appears at the start of the book, and is called, ‘The current state of the art.’ We also decided to adhere to geographers’ current recommendation to use the term ‘south-west Asia’ instead of ‘Middle East’ and ‘Near East,’ which are both ambiguous and Eurocentric geographic terms.

Acknowledgements We wish to express our gratitude to the many colleagues who helped us in the preparation of this edition. We are particularly indebted to those who advised us on the current state of research in their regions; this was essential in our ability to present a much revised edition. Among them are: Felix Bittmann, Ksenija Borojevic, Laurent Bouby, Otto Brinkkemper, Ramon Buxó, Ahmed Fahmy, Andrew Fairbairn, Ferenc Gyulai, Maria Hajnalova, Roman Hovsepyan, Stefanie Jacomet, Glynis Jones, Sabine Karg, Mordechai Kislev, Marianne

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PREFACE TO THE FOURTH EDITION

Kohler-Schneider, Angela Kreuz, Terttu Lempiäinen, Elena Marinova, Felicia Monah, Dominique de Moulins, Aldona Mueller-Bieniek, Mark Nesbitt, Simone Riehl, David Earle Robinson, Mauro Rottoli, Anaya Sarpaki, Margareta Tengberg, João Tereso, Tania Valamoti, Marijke van der Veen, Karin Viklund, George Willcox. Mordechai Kislev, Moshe Feldman, and particularly Mark Nesbitt, offered valuable criticism in reading an initial revised draft. We are grateful to Ori Fragman, Mordechai Kislev, Linda Learn (Class Act Fabrics), Oxford University Press, for providing new pictures for this edition, to Reuven Soffer from Soffer Cartography inc., Jerusalem, for producing all maps, and to the Photography and Graphic Design Departments, Weizmann Institute of Science, for

the new and renewed pictures and tables. Also, to Dorian Fuller for his permission to photograph Aegilops tauschii from his reference collection, and to Sue Colledge for sharing with us her Access database of archaeobotanical finds across southwest Asia and Europe. Special thanks are due to Elisabetta Boaretto for her assistance with updating radiocarbon dating, and to Aaron Rottenberg for his assistance with updating the molecular analysis. We also want to thank Anat Hartmann-Shenkman, Yael MahlerSlasky, Leon Cherniaev and Ravit Ferera for their assistance in preparing this edition. D.Z.

Jerusalem

E.W.

Ramat-Gan & Rehovot 2011

Preface to the third edition

In the last seven years, considerable progress has been made in our understanding of the origin and spread of cultivated plants in west and central Asia, in the Mediterranean basin, and in the temperate parts of Europe. Today the wild ancestors of the crops that initiated and sustained food production in this part of the world are already well-identified. Moreover, the archaeobotanical evidence assembled on the origins and spread of these cultivated plants is today much more extensive and convincing than the data available only a few years ago. In this third edition, an attempt is being made to integrate the new evidence and to update the book. Significantly, the additional information does not contradict the main conclusions presented in the first (1988) and second edition (1993). Moreover, it adds considerably to the clarity of the portrayed picture. To keep the book more or less to its original size, we decided to focus on domestication, and to omit

the chapter on fruit collected from the wild that appeared in earlier editions. The knowledge about collection from the wild has increased greatly over the last few years; we believe the subject deserves to be treated separately. We wish to express our gratitude to the many colleagues who helped us in the preparation of this edition. We are particularly indebted to Corrie Bakels, Dorian Q. Fuller, David R. Harris, Gordon C. Hillman, Stefanie Jacomet, Mordechai E. Kislev. Karl-Heinz Knötrzer, Helmut Kroll, Desanka Kučan, Naomi F. Miller, Mark Nesbit, Jurgen SchultzeMotel, Krystyna Wasyliltowa. George H. Willcox and Willem van Zeist. D.Z. M.H.

Jerusalem Mainz 2000

Preface to the second edition

Since the completion of the writing of the first edition in 1987, archaeobotanical investigations and the study of Old World crops and their wild relatives has continued apace. An impressive body of new evidence about crops and sites was added. Significantly, the new information does not contradict the main conclusions that were drawn five or six years ago. It confirms them. In this second edition, an attempt is made to integrate this new evidence. We have also filled a gap by adding a chapter on dye plants. The revision is

most apparent in the vegetables, in the fruit trees, and in some of the minor grain crops. Only a few years ago, our knowledge of their origin and early history was embarrassingly fragmentary. At least for some of these crops, the evidence today permits a sounder synthesis. D.Z. M.H.

Jerusalem Mainz 1993

Preface to the first edition

South-west Asia, Europe, and the Nile valley arc unique today for the vast extent of archaeobotanical exploration. In the last thirty years, hundreds of Mesolithic, Neolithic, and Bronze Age sites have been excavated in these territories. Plant remains in many sites have been expertly identified, culturally associated, and radiocarbon-dated, and the finds have offered critical information on the plants that formed the start of agriculture in this part of the world. Considerable progress has also been achieved in the field of the wild ancestry of Old World crop. The wild progenitors of most of these cultivated plants have now been satisfactorily identified, both by comparative morphology and by genetic analyses. The distribution and ecological ranges of the wild relatives have been established, and furthermore, comparisons between wild types and their cultivated counterparts have revealed the evolutionary changes which were brought about by domestication. As a result of these achievements, south-west Asia, Europe, and Egypt emerge as the first major geographical area in the world in which the combined evidence from archaeology and living plants

permits a modern synthesis of crop-plant evolution. The accumulated information provides reasonable answers to the following questions: (a) What were the first plants to be domesticated in the Old World? (b) Where can the earliest signs of their domestication be found? (c) What were the subsequent main developments in plant cultivation over these regions? (d) What crops were introduced into this area from other parts of Asia and Africa? (c) When did all these events take place? In the following chapters an attempt is made to answer these questions and provide a review of the origin and the spread of cultivated plants in southwest Asia, Europe, and Africa north of the Sahara, i.e. the classical ‘Old World’. The aim was to trace plant domestication and crop-plant evolution in this part of the globe from its early beginnings up to classical times. The treatment (Chapters 2–9) is crop by crop. Chapter 10 adds essential documentation on representative archaeological sites. The information given is based on work published up to 1985. D.Z. M.H.

Jerusalem Mainz 1987

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Contents

1 Current state of the art Beginnings of domestication Neolithic south-west Asian crop assemblage Wild progenitors The spread of south-west Asian crops Availability of archaeological evidence Early domestication outside the ‘core area’ Beginning and spread of horticulture Vegetables Weeds and crops Migrants from other agricultural regions

1 1 1 3 4 4 5 5 6 7 7

2 Sources of evidence for the origin and spread of domesticated plants Archaeological evidence Evidence from the living plants Radiocarbon dating and dendrochronology

9 9 13 17

3 Cereals Wheats: Triticum Einkorn wheat: Triticum monococcum Emmer and durum-type wheats: Triticum turgidum Bread wheat: Triticum aestivum Timopheev’s wheat: Triticum timopheevii Barley: Hordeum vulgare Rye: Secale cereale Common oat: Avena sativa Broomcorn millet: Panicum miliaceum Foxtail millet: Setaria italica Latecomers: sorghum and rice

20 23 34 39 47 51 51 59 66 69 71 72

4 Pulses Lentil: Lens culinaris Pea: Pisum sativum Chickpea: Cicer arietinum Faba bean: Vicia faba Bitter vetch: Vicia ervilia

75 77 82 87 89 92 xiii

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CONTENTS

Common vetch: Vicia sativa Grass pea: Lathyrus sativus Spanish vechling: Lathyrus clymenum Fenugreek: Trigonella foenum-graecum Lupins: Lupinus

95 95 97 97 98

5 Oil- and fibre-producing crops Flax: Linum usitatissimum Hemp: Cannabis sativa Old World cottons: Gossypium arboreum and G. herbaceum Poppy: Papaver somniferum Gold of pleasure: Camelina sativa Other cruciferous oil crops Sesame: Sesamum indicum

100 101 106 107 109 111 112 112

6 Fruit trees and nuts Olive: Olea europaea Grapevine: Vitis vinifera Fig: Ficus carica Sycamore fig: Ficus sycomorus Date palm: Phoenix dactylifera Pomegranate: Punica granatum Apple: Malus domestica Pear: Pyrus communis Plum: Prunus domestica Cherries Prunus avium and P. cerasus Latecomers: apricot, peach, and quince Carob: Ceratonia siliqua Citrus fruits Almond: Amygdalus communis Walnut: Juglans regia Chestnut: Castanea sativa Hazelnut: Corylus avellana Pistachio: Pistacia vera

114 116 121 126 130 131 134 135 138 140 143 144 145 146 147 149 150 151 151

7 Vegetables and tubers Watermelon: Citrullus lanatus Melon Cucumis melo Leek: Allium porrum Garlic: Allium sativum Onion: Allium cepa Lettuce: Lactuca sativa Chufa or rush nut: Cyperus esculentus Cabbage: Brassica oleracea Turnip: Brassica rapa Beet: Beta vulgaris Carrot: Daucus carota Celery: Apium graveolens

153 153 154 155 156 157 157 158 158 159 159 160 160

CONTENTS

Parsnip: Pastinaca sativa Asparagus: Asparagus officinalis

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161 161

8 Condiments Coriander: Coriandrum sativum Cumin and dill: Cuminum cyminum and Anethum graveolens Black cumin: Nigella sativa Saffron: Crocus sativus

163 163 164 164 165

9 Dye crops Woad: lsatis tinctoria Dyer’s rocket: Reseda luteola Madder: Rubia tinctorum True indigo: Indigofera tinctoria Safflower: Carthamus tinctorius

166 166 167 167 168 168

10 Plant remains in representative archaeological sites Iran Iraq Turkey Syria Israel and Jordan Egypt Libya Morocco Caucasia and Transcaucasia Central Asia Cyprus Greece Crete Former Yugoslavia Bulgaria Rumania Moldavia and Ukraine Hungary Austria Italy Poland Czech Republic and Slovakia Switzerland Germany The Netherlands Belgium Denmark Sweden Norway Finland Britain and Ireland

169 169 170 171 172 173 174 176 176 176 176 177 177 179 179 179 181 181 182 183 184 185 185 186 188 189 189 189 190 190 190 190

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CONTENTS

France Spain Portugal

191 192 193

Appendix A: Site orientation maps

194

Appendix B: Chronological chart for the main geographical regions mentioned in the book

197

Appendix C: Information on archaeological sites which appear on Map 2

200

References

201

Index

237

C H A PTER 1

Current state of the art

The aim of this book is to review available information on the origin and spread of domesticated plants in south-west Asia, Europe, and the Mediterranean Basin. Two sources of evidence exist: firstly, information obtained by the analysis of plant remains retrieved from archaeological excavations, where early archaeological contexts—namely Epipalaeolithic/Mesolithic, Neolithic, and Bronze Age cultures—are the main source; and secondly, data provided by living plants, particularly by the wild progenitors of domesticated plants. This chapter presents the conclusions of the book as determined from the combined information provided by these two sources (relevant data and references will be presented in the following chapters).

Beginnings of domestication The first definite signs of domesticated plants in the Old World appear in a string of Early Pre-Pottery Neolithic B (PPNB) farming villages that developed in south-west Asia (Map 1) by ca. 10,500–10,100 calibrated years before present (cal BP). Spikelet forks of emmer and einkorn wheat with telltale, rough disarticulation scars (pp. 24, 30–31) provide the most convincing evidence that these cereals were already domesticated by this time, and in this area. The contemporary appearance of relatively plump kernels further supports this notion, but cannot be regarded as a fully reliable indication of the early stage of domestication. These remains and further evidence of pre-domestication cultivation suggest that the actual beginning of wheat cultivation in this area should have been even earlier. No convincing pre-PPNB domesticated plants have yet been found.

There is a scholarly debate as to whether agriculture originated in several places across a wide area, including the Levant and northern Fertile Crescent (e.g. Weiss et al. 2006; Willcox et al. 2008), or whether it evolved in only one part of the Fertile Crescent, such as south-east Turkey (e.g. Lev-Yadun et al. 2000). Although current archaeobotanical data support the first view, this critical question requires more archaeobotanical and radiocarbon dating evidence to support any definitive finding.

Neolithic south-west Asian crop assemblage The crops of early Neolithic agriculture in southwest Asia are fairly well recognized. The most numerous vegetable remains in early farming villages come from three cereals: emmer wheat (Triticum turgidum subsp. dicoccum), einkorn wheat (T. monococcum subsp. monococcum), and barley (Hordeum vulgare). Diagnostic morphological traits (non-brittle ears, broad kernels) traceable in the archaeological finds indicate that by 10,500–10,100 cal BP, these domesticated annual grasses were intentionally sown and harvested in a string of PrePottery Neolithic B sites in south-west Asia. Emmer wheat and barley seem to have been the more common crops. Einkorn wheat is somewhat less apparent. Several grain legumes appear as constant companions of the cereals (see Map 2—Plate 6). The most frequent pulses in the early Neolithic south-west Asian contexts are lentil (Lens culinaris) and pea (Pisum sativum). Two more local legume crops are bitter vetch (Vicia ervilia) and chickpea (Cicer arietinum). In contrast to the cereals, archaeological

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DOMESTICATION OF PLANTS IN THE OLD WORLD

Asikli Höyük

Çayönü Cafer Höyük

10,500-10,000 BP 10,000-9,500 BP Tell Abu Hureyra

9,500-9,000 BP

einkorn wheat emmer wheat

Jarmo

Kissonerga-Mylouthkia + Shillourokambos Yiftah'el

barley chickpea

Jericho

Tell Aswad

‘Ain Ghazal

flax

Ali Kosh

lentil pea bitter vetch 0 0

100 200

200 miles 400km

Map 1 Archaeological sites in which the earliest south-west Asian domesticated grain crops were reliably identified.

remains of pulses usually lack morphological features by which initial stages of domestication can be recognized. Clear indications of lentil domestication appear at about 10,100–9,700 cal BP; and of pea, chickpea, and bitter vetch, at about 9,900–9,500 cal BP. Probably all four legumes were cultivated somewhat earlier, either together with wheats and barley or soon after the domestication of those cereals. Finally, flax (Linum usitatissimum) belongs to the south-west Asian group of founder crops. It is impossible to decide whether the material obtained from Early Neolithic layers represents collected wild flax or the remains of domesticated forms. Yet, as in the case of the legumes both direct and circumstantial evidence indicates that by 9,900–9,500 cal BP, flax was already domesticated in south-west Asia. Evidence for early domestication of additional plants in south-west Asia is much less convincing. Grass pea (Lathyrus sativus) might have been such a crop, yet the bulk of its early remains comes from

eighth and seventh millennia BP sites in Greece and Bulgaria. Signs that rye (Secale cereale) was a southwest Asian Neolithic crop are much rarer. The origin and early spread of the faba bean (Vicia faba) is even less clear. The plant remains from south-west Asian PrePottery Neolithic B (PPNB) sites reveal another feature: as a rule, not a single crop but rather a combination of cereals, pulses, and flax appears in these early farming villages. Moreover, the assemblage seems to be similar throughout the Fertile Crescent (see Map 2—Plate 6). In other words, a common package of grain crops characterizes the development of agriculture in this ‘core area’. At almost the same time, signs of herding appear, implying that sheep and goats had also been brought under human control. Shortly after, cattle and pig domestication took place (Zeder 2011). Thus, an effective south-west Asian Neolithic food-production ‘package’ was formed, comprising

CURRENT STATE OF THE ART

60

Legened

61

einkorn wheat emmer wheat barley flax lentil pea

58 62

71

2,500-2,000 BP 3,000-2,500 BP 3,500-3,000 BP 4,000-3,500 BP 4,500-4,000 BP 5,000-4,500 BP 5,500-5,000 BP 6,000-5,500 BP 6,500-6,000 BP 7,000-6,500 BP 7,500-7,000 BP 8,000-7,500 BP 8,500-8,000 BP 9,000-8,500 BP 9,500-9,000 BP 10,000-9,500 BP 10,500-10,000 BP

63 59

72

70

66

69

65 68

40

56

67

53

57

39

52,55

73

48

21 37 36

47

49

50 82

38

35 54

51

75

78

64

41

22

33

34 44

81

3

1

23

32

4

74

25 6 31

77

24

29

42 43

79

7

27 45

76

5

28 30

26

18 8

19

80

3

46

9

17

2

14

20

10 13 11 12 16

0 0

Scale 1:16,000,000 125 250 375 250

15

500 miles

500 km 83

Map 2 The spread of the south-west Asian Neolithic crop assemblage in Europe, west Asia, and north Africa. For details on the numbered sites, see Appendix C (p. 200). These are the earliest sites in which domesticated grain crops were found, in each country. (See Plate 6.)

vegetative crops as well as domestic animals. Indeed, the remains uncovered in south-west Asian PPNB sites indicate a major shift in food practices. While in Epi-Palaeolithic contexts, gathering and hunting of a wide spectrum of wild species is apparent, the PPNB farmers already appear to focus on domesticates as their principal source of food. A large proportion of the remains retrieved from these early farming sites belong to the crops mentioned above and domestic animals. There is also a sharp quantitative and qualitative drop in the wild-species intake. An important confirmation of this ‘package’ concept occurred recently with the discovery of just such an ensemble of plants and animals in Early PPNB Cyprus, although some of them were not yet strictly domesticated.

Wild progenitors The wild ancestors of most of the food plants of south-west Asia, Europe, and the Mediterranean

Basin are already well identified. The distribution areas and the main ecological preferences of most of them are also well known. Comparison of this evidence with the archaeological findings reveals that with practically all early crops, the first signs of domestication appear in the same general areas where the wild ancestral stocks abound today. The geographic distribution of the wild progenitors of Neolithic grain crops is significant. Apart from flax and barley, the wild ancestors of the founder crops have a rather limited distribution. Wild emmer wheat and wild chickpea are endemic to the Fertile Crescent. Assuming that their distribution did not change drastically during the last ten millennia, the domestication of these crops could only have taken place in this restricted area. Because domesticated emmer wheat appears to be the most important Neolithic crop throughout south-west Asia, Europe, and the Mediterranean Basin, the confinement of its wild progenitor to the Fertile Crescent delimits the place of origin of this

4

DOMESTICATION OF PLANTS IN THE OLD WORLD

domesticated cereal. It also marks the rather restricted geographic area where Old World Neolithic agriculture could have originated. Wild forms of einkorn wheat, lentil, pea, and bitter vetch have a somewhat wider distribution, but all, including barley, are centered in the Fertile Crescent; that is, the region in which the earliest farming villages have been discovered.

The spread of south-west Asian crops A most remarkable feature of south-west Asian Neolithic agriculture is its rapid expansion soon after establishment in the nuclear area (see Map 2—Plate 6). The quality and quantity of available archaeobotanical evidence varies considerably from region to region. Comprehensive information is available for most parts of Europe, but there is much sparser and frequently incomplete documentation from Caucasia, Eastern Europe, and central Asia. In Africa, critical data on plant remains are available only for Egypt (but a few current projects might add vital data for north Africa). In spite of the uneven documentation, the following main features of the diffusion of agriculture seem apparent. The spread of agriculture from its south-west Asian core to Europe and central Asia involves the species contained in the Neolithic crop assemblage. Map 2 (Plate 6) summarizes the information about the six most important south-west Asian crops: emmer wheat (including its free-threshing derivatives), einkorn wheat, barley, lentil, pea, and flax. From the data presented in this map and in Chapter 10, it is evident that crops domesticated in the south-west Asian core area were the initiators of food production in Europe, central Asia, and the Mediterranean Basin (including the Nile Valley). The earliest farming cultures in these vast regions always contain wheat and barley, with one, two, or more of the other south-west Asian founder crops frequently present as well. Establishment of the south-west Asian crop assemblage in the Fertile Crescent and its spread both west (to Europe) and east (to central Asia and to the Indian subcontinent) was rapid (see Map 2—Plate 6). From the first farming communities in the ‘Levantine Corridor’ at ca. 10,500–10,200 cal BP, it was found to cover the whole Fertile Crescent by

9,500–9,000 cal BP. By ca. 9,000–8,500 cal BP, agriculture had already appeared in Crete and Greece. By the end of the ninth millennium BP, these crops were grown in Obre in Bosnia-Hercegovina and in Jeitun in Turkmenia. Soon after, agriculture appears as far west as Balma Margineda in Andorra, Spain, and Sacarovca in Moldavia—and as far south as Grotta dell’Uzzo in Sicily. By the second half of the eighth millennium BP, the Linearbandkeramik farming culture was already firmly established in loess soil regions throughout central Europe, extending to Poland in the east, to northern France, and Germany in the west. At the same time, early Neolithic farming villages appeared in south Spain, the Nile Valley, and in Chokh in Caucasia. Substantial information on the age and spread of early farming cultures is available for Europe, where radiocarbon dating of sites exhibiting evidence of early farming enabled the reconstruction of the diffusion of agriculture. The evidence from Caucasia, central Asia, and eastern Europe is much more fragmentary. Yet the finds retrieved from sites including Jeitun (p. 176) demonstrate that the diffusion of the south-west Asian crops towards central Asia happened relatively early, although it took longer to reach Transcaucasia and the Nile Valley. All over these vast areas, the start of food production involved the same south-west Asian crops.

Availability of archaeological evidence Any attempt to reconstruct the origins and diffusion of agriculture in Eurasia and Africa must address the uneven archaeological record. As already mentioned, plant remains of Europe, south-west Asia, and the Mediterranean Basin provide us with a reasonable overview of the beginnings and development of agriculture in these major areas. In contrast, the archaeobotanical evidence from central and eastern parts of Asia and from eastern Europe is much less complete. It is very poor in Africa north of the Sahara. Consequently, while the early stages of food production in south-west Asia are relatively well documented, most founder crops are adequately identified, and the expansion to Europe and west Asia are convincingly elucidated, there are far fewer solid facts on crop domestication and the development of farming in east Asia (Smith

CURRENT STATE OF THE ART

1998). However in the last few years, archaeobotanical findings in these agricultural domains have improved considerably. The history of crop domestication in the African Savanna belt is still largely uncharted and we still know very little about the evolution of the unique crop assemblage of this region (Harlan 1992a). The time and place of origin of the majority of the east and south Asian crops, and of practically all the sub-Saharan African crops, are yet not fully established. In numerous cases, the wild progenitors have not yet been satisfactorily identified or they are only very superficially known. However, critical archaeobotanical information has been assembled on at least two principal crops; rice (Oryza sativa) and foxtail millet (Setaria italica). Their essential role in the independent rise of farming in China is now well documented. At present, our picture of crop-plant evolution in Eurasia and Africa is unbalanced. While there is relatively reliable information on its development in the classical Old World, we are largely uninformed of events south and east of this area. We also know relatively little about the early interactions between west Asia and the major agricultural provinces in east and south Asia, and in Africa south of the Sahara.

Early domestication outside the ‘core area’ Signs of additional domesticants start appearing soon after the introduction of south-west Asia agriculture to Europe, central Asia, and the Mediterranean Basin. Addition of some of these crops obviously took place outside south-west Asia, but they developed within the already established agriculture of the south-west Asian crop assemblage. The poppy, Papaver somniferum, provides a well-documented example of such domestication. Both the area of distribution of the wild poppy and the archaeological finds (p. 109–111) indicate that P. somniferum was brought into domestication in west Europe. It was added to the south-west Asian grain-crop assemblage after the latter’s establishment in western Europe. Chufa, Cyperus esculentus, is another example of an early local addition, this time in the Nile Valley (p. 158).

5

Its dry tubers were found in large quantities in Egypt from pre-dynastic times on. The early appearance of broomcorn millet, Panicum miliaceum, in the Caspian basin and the Czech Republic (pp. 69–70) might indicate another local addition. However, since the archaeological evidence from central and east Asia is still inadequate, it is impossible to decide whether the Caspian P. miliaceum was added to the expanding south-west Asian crop assemblage after it reached central Asia, or whether this cereal represents an east Asiatic domestication independent of the southwest Asian diffusion.

Beginning and spread of horticulture Olive, grapevine, fig, and date palm seem to have been the first principal fruit crops domesticated in the Old World. Definite signs of olive and date-palm domestication appear in Chalcolithic Levant about 6,800–6,300 cal BP. Indications of date-palm domestication are also available from contemporary lower Mesopotamia. We still do not know the extent of Chalcolithic horticulture. Except for the IsraelJordan area, the archaeobotanical information from seventh–sixth millennia BP sites in the Levant is still insufficient. The picture changes drastically in the Early Bronze Age (first half of the fifth millennium BP). From this time on, olives, grapes, and figs emerge as important additions to grain agriculture, initially in the Levant and soon after, in Greece. These crops were subsequently planted throughout the Mediterranean Basin. The extensive Bronze Age cultivation of olives and grapes is indicated by the appearance of numerous presses and remains of storage facilities for olive oil and wine. At the same time, dates were domesticated on the southern fringes and the warm river basins of the south-west Asia, and they abound in the Nile Valley during the New Kingdom. Apple, pear, plum, and cherry seem to have been added much later to Old World horticulture, as definite signs of their domestication appear only in the first millennium BC. Their culture is almost entirely based on grafting, so they could have been domesticated extensively only after the introduction of this sophisticated method of vegetative propagation.

6

DOMESTICATION OF PLANTS IN THE OLD WORLD

Remains of fruit trees rarely show diagnostic anatomical traits enabling archaeobotanists to distinguish between fruits collected from the wild or those harvested from domesticated orchards. To a large extent, recognizing domestication in fruit crops is based on circumstantial evidence, such as the finding of fruit remains in areas in which the wild forms do not occur or on the quantitative analysis of artefacts associated with fruit products (e.g. oil, wine). It is difficult, therefore, to determine the initial stage of fruit crop domestication: in other words, it might well be that olive, grape, fig, or date cultivation did not originate in the Chalcolithic (sixth millennium BP), but was already active in the late Neolithic (seventh millennium BP). Despite these uncertainties, the following have been confirmed: (a) the earliest definite signs of fruit tree domestication appear in the south-west Asia; (b) horticulture developed only after the firm establishment of grain agriculture; (c) as with grain crops, several local wild fruits were taken into domestication at about the same time; (d) domestication of fruit crops relied heavily on the invention of vegetative propagation; (e) planting of perennial fruit trees is a long-term investment, promoting a fully settled way of life; (f) soon after its successful establishment, horticulture spread from its original ‘core area’ into new territories in the Mediterranean Basin and south-west Asia; and (g) after the introduction of grafting (pp. 114–115), the domestication of a whole group of ‘second-wave’ fruit crops became possible. Available archaeobotanical evidence of the beginning of fruit-crop domestication can also be supported by information on the wild relatives. Wild olive, grapevine, fig, and date are widely distributed over the Mediterranean and southwest Asia. They have a wide geographic distribution, so this by itself does not provide critical values for a precise delimitation of the place of origin of these fruit crops. Yet it is reassuring to know that forms from which domesticated clones could have been derived thrive in wild niches in the east Mediterranean basin. Therefore, evidence from the living plants complements the archaeological finds. Most probably olive, grapevine, date, fig, as well as pomegranate and almond, were first brought into domestication in the same

general area where, several millennia earlier, grain agriculture was successfully established in the Old World. Thus, during the sixth millennium BP, eastern Mediterranean Basin human societies belonging to the Chalcolithic and Bronze Age cultures, were introduced to the use of copper and bronze, and they also mastered horticulture.

Vegetables This is the least-known group of domesticated food plants of the Old World. Vegetable material consists almost entirely of perishable soft tissues, which stand a meagre chance of charring and surviving as archaeological remnants (p. 153). Consequently, only few vegetable remains have been detected in excavations. The exceptions here are Egyptian and Judean Desert caves. In Egypt, especially arid country vegetables placed in pyramids and graves commonly survived by desiccation, and show that garlic, leek, onion, lettuce, melon, watermelon, and chufa were cultivated in the Nile Valley in the second and the first millennia BC. As amply described by Keimer (1924, 1984), vegetable gardens constituted an important element of food production in Egyptian dynastic times. Beyond Egypt there are almost no early archaeobotanical finds of vegetable crops. However, early literary sources show that by the start of the second millennium BC, vegetable gardens flourished not only in the Nile Valley but also in Mesopotamia. Furthermore, in both areas the crops grown were more or less the same. The only major exception was chufa which was restricted, almost entirely, to Egypt. In summary, available evidence makes it clear that by the Bronze Age vegetable crops were part of food production both in Lower Mesopotamia and in Egypt. It is very likely that this geographic pattern is not accidental. In both regions, we are faced with the dense human settlement of very arid environments. Survival in these zones depends on utilization of limited areas of irrigated or flooded land which is bordered by large, barren deserts. Areas with no vegetation have little to offer in the way of supplementary resources of green wild plants. This shortage invites human initiative. The early development of vegetable gardens might have been

CURRENT STATE OF THE ART

caused by such needs. It must be taken into consideration that this picture is partly skewed by the lack of evidence in other regions.

Weeds and crops Several Old World grain plants, oil producers, and vegetables seem to be ‘secondary crops’; that is, they first evolved as weeds and were only later established as crops (p. 16). Oat, Avena sativa, rye, Secale cereale subsp. cereale, and gold of pleasure, Camelina sativa, are well-documented examples of this mode of evolution under domestication. Turnip, lettuce, carrot, beet, leek, and several other vegetables are also very likely to have entered domestication through the same ‘back door’. The incorporation of secondary crops into Old World food production seems to have happened rather late, since definite signs of their domestication appear in Europe and west Asia only in the second and first millennia BC.

Migrants from other agricultural regions With few exceptions, the classical ‘Old World’ (south-west Asia, the Mediterranean Basin, and temperate Europe) received crops from other agricultural regions rather late in its agricultural history. Foreign crops that arrived in this area (in pre-Columbian times) fall into the following geographical groups (Zohary 1998):

(a) Temperate climate crops from central and/or east Asia Broomcorn millet (Panicum miliaceum) and foxtail millet (Setaria italica) seem to represent the earliest arrivals. The origin of P. miliaceum is not fully understood, but it was probably taken into domestication in central Asia–north China (p. 69–71). It already appears in Caucasia and in central Europe in sites around the first half of the eighth millennium BP. S. italica, now recognized as a founder crop of north China agriculture (p. 71), appeared in central Europe in the first half of fourth millennium BP, some four thousand years later. For millet, as well, the available information suggests arrival from the east (p. 69). However, the pos-

7

sibility of independent domestication of foxtail millet in the west has not been ruled out yet. Hemp (Cannabis sativa) reached Anatolia and Europe much later. Its remains appear (pp. 106–107) from the eighth century BC onwards. Apricot (Armeniaca vulgaris) and peach (Persica vulgaris) could have been taken into domestication either in central Asia or in China (pp. 154–155); the domesticated pistachio (Pistacia vera) must have originated in central Asia (pp. 151–152). The peach seems to have reached the Mediterranean Basin by the middle of the first millennium BC. Apricot and pistachio arrived only in Roman times.

(b) Warm-weather crops from south and/or east Asia A group of more tropical crops (sensitive to freezing temperatures) that originated in south and/or east Asia, seem to have migrated into the south-west Asia and the Mediterranean Basin from the Indian subcontinent. Many of these cultigens were already grown in India and Pakistan in the second millennium BC. Sesame (Sesamum indicum) is apparently the earliest of these migrants (pp. 112–113). Undisputed remains of this Indian oil crop already appear in south-west Asia in Iron Age (ca. 900–600 BC) contexts. The citron (Citrus medica) was grown in the east Mediterranean basin (p. 146) by the end of the fourth century BC. Asian rice (Oryza sativa) seems to have arrived (pp. 73–74) in Hellenistic or early Roman times. The cucumber (Cucumis sativus) might also have been introduced at the same time (p. 155). Finally, Old World cottons (Gossypium arboreum and/or G. herbaceum) could have already spread from the Indian subcontinent into the southwest Asia (pp. 107–109) during Roman rule. However, a fully developed cotton industry appeared in this area only in Early Islamic times. An impressive introduction of Indian and southeast Asian crops was undertaken by the Arabs soon after their conquests (Watson 1983; Zohary 1998). The Early Islamic diffusion (eigth–eleventh centuries AD) includes lemon (Citrus limon), lime (C. aurantiifolia), bitter orange (C. aurantium), pummelo (C. maxima), and indigo (Indigofera tinctoria)— all of which are discussed in this book. It also involves sugar cane (Saccharum officinarum) and

8

DOMESTICATION OF PLANTS IN THE OLD WORLD

sugar extraction technology, banana and plantain (Musa cultivars), aubergine (Solanum melongena), and taro (Colocasia esculenta (L.) Schott), although these crops are not surveyed here.

(c) Warm-weather crops from Africa south of the Sahara Although there are several good reasons to assume (Harlan 1992) that indigenous agriculture was already well developed in sub-Saharan Africa by 1,000 BC (or even earlier), surprisingly few of the native African cultigens spread north into the Mediterranean Basin. This is even more puzzling since several African grain crops, namely sorghum

(Sorghum bicolor), pearl millet (Pennisetum glaucum), and cow pea (Vigna unguiculata), seem to have reached the Indian subcontinent already by the second millennium BC (Possehl 1998; Fuller 2000; Manning et al. 2011). In contrast, only few arrivals are recorded north of the Sahara. Domesticated sorghum, was grown in Egyptian Nubia from ca 100 AD onwards (pp. 72–73), yet there are no signs of its spread further north. Advanced durra-type sorghum cultivars appear in south-west Asia only in Early Islamic times, and as Harlan and Stemler (1978) argue, they might have arrived not from Africa but from India. In addition, cowpea (Vigna unguiculata) is known to have come from Egypt in Hellenistic and Roman times (Germer 1985, p. 88).

C H A PTER 2

Sources of evidence for the origin and spread of domesticated plants

The study of the origin and spread of domesticated plants is an interdisciplinary venture based on evidence from numerous sources. Several disciplines, such as archaeology, botany, genetics, chemistry, anthropology, agronomy, and linguistics are involved (for review, see Harlan and de Wet 1973), yet the different sources of evidence vary considerably in reliability and relative weight. The modern synthesis leans heavily on two principal sources:

Readers who wish to acquaint themselves with some of the more recent research should consult Pearsall (2000) for archaeobotany, and Hancock (2004) for plant evolution, genetics, and crop sciences. All serve as a good sources for initial orientation. The following sections review the main sources of evidence on which the modern assessment of cropplant evolution is based. A list is given in Table 1.

(i) information obtained through examination of plant remains retrieved from archaeological excavations; and (ii) evidence gathered from living plants, particularly the main traits that evolved under domestication (‘domestication syndrome’) and the genetic affinities between the crops and their wild relatives.

Archaeological evidence

Since 1950s major discoveries have radically changed our view on the origin of domesticated plants. They have transformed a realm of plentiful speculation and few solid facts into a welldocumented field. In the light of this, the contributions of several classical tools had to be fundamentaly reconsidered. Some sources of evidence, such as N.I. Vavilov’s monumental studies of variation and distribution of crops (Vavilov 1949–50; 1987), had to be considerably revised and re-evaluated. Others, such as linguistic comparisons (widely used by de Candolle 1886), appear to have retained some relevance, even today. However, with the flood of archaeological documentation, genetic information, and new techniques from molecular science, those discoveries now carry less weight.

(i) When and where do we find the earliest signs of domesticated crops? (ii) How and when did the crops spread to attain their present distributions? (iii) What were the early cultigens like? (iv) What were the main changes in the crops once they were introduced into cultivation? (v) Where and when did these changes take place?

The primary contribution of archaeology to the understanding of crop-plant evolution is by the recovery of plant remains in archaeological excavations, and by identifying to what crop species they belong. The accumulated evidence contributes to answering the following questions:

Obviously the key to all answers is the availability of ‘fossil evidence’; that is, sufficient amounts of dated, culturally defined plant remains amenable to analysis. The following sections survey the main conditions under which plant material survives in archaeological contexts. These are also listed in Table 2. 9

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DOMESTICATION OF PLANTS IN THE OLD WORLD

Table 1 Sources of evidence on the origin and spread of domesticated plants I. Archaeological evidence 1. Archaeobotany: Identification of plant remains retrieved from archaeological excavations in connection with cultural associations and 14 C-dating. Determination of the earliest signs of domestication in these plants and their subsequent spread. Changes in crops in time and space. Crop assemblages in various cultures. 2. Additional evidence (a) Artefacts. Evaluation of: (i) dated tools associated with cultivation, harvesting, and processing of crops; (ii) cultivation artefacts such as irrigation canals, terraces, lynchets, plough marks, and cultivation boundaries. (b) Art. Early drawings, paintings, and reliefs of domesticated plants. (c) Palynology. Appearance of pollen grains of crops and weeds in dated cores or site contexts. (d) Weeds associated with agriculture. (e) Examination of ancient DNA extracted from plant remains. (f) Chemical analysis. Identification of crops by specific organic residues retained in charred seeds, ancient vessels, charcoal, etc. (g) Starch analysis. Identification of plant remains and usage of tools by the remains of starch granules. II. Evidence from the living plants 1. Search for the wild progenitors. Identification of the nearest wild relatives of the domesticated crops by use of: (a) comparative morphology and comparative anatomy (classical taxonomy). (b) determination of genetic affinities by cytogenetic analysis. (c) determination of genetic affinities by DNA and protein resemblances. 2. Distribution and ecology of the wild progenitors (a) Geographic distribution of the wild relatives (including weedy forms). (b) Characterization of the habitats and the main adaptations of the wild relatives. 3. Evolution under domestication Main trends of morphological, physiological and chemical changes. The range and the structuring of genetic variation in the crops and in their wild progenitors. Development of crop complexes (wild forms, weedy races, and cultigens). Methods of planting, maintenance, and usage. 4. Additional evidence (a) Genetic systems: characterization of the main systems operating under domestication, especially reproductive systems (including vegetative propagation). (b) Genetic interconnections between cultivars and wild relatives. (c) Intentional and unconscious selections. III. Other pertinent sources 1. Historical information. Representation of the plants in art, documentation in inscriptions, tablets, manuscripts, and books. 2. Linguistic comparisons. Names of crops in various languages. 3. Circumstantial evidence: Geological, climatic, hydrological, limnological, dendrochronological, anthropological, and zoological indications on the initiation and spread of agriculture.

Charred remains Charred (carbonized) remains are the commonest source of plant material in archaeological excavations that are available for analysis. Carbonization occurs on exposure to high temperatures, in most cases due to fires. Such heating (under a limited supply of oxygen) converts the plant’s organic compounds into charcoal. Since bacteria, fungi, or other decomposing organisms do not affect charcoal, carbonized plant remains survive in most environments. This includes wet places where ordinary organic

material decays rapidly. Carbonized plant remains in archaeological contexts are therefore not products of geological carbonization (true fossils). They represent only ‘subfossil’ elements charred by fire. When slowly and mildly charred, wood, seeds, nuts, and sometimes even fleshy fruits, parenchymatous storage tissues, or ears of cereals, can still retain most of their morphological and anatomical features. The morphology and the microscopic anatomical structures are frequently preserved in astonishing clarity. This allows a reliable identification of the plant remains.

SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS

Table 2 Preservation of plant remains in archaeological excavations I. Charred remains 1. Charred during handling: (a) near a hearth/oven (b) in a drying kiln (c) in a storage pit/silo (when cleaned) (d) in pottery fired in a pottery kiln 2. Charred by conflagration: (a) stored material (b) material embedded in daub, unfired bricks, and floors (c) thatching material (d) scattered or dumped material II. Plant impressions 1. In pottery 2. In bricks and daub III. Parched remains 1. In arid regions: (a) in caves (b) in tombs and pyramids (c) in clay 2. In temperate regions: (a) in sealed containers (b) in offerings embedded in walls IV. Waterlogged remains 1. In lakes 2. In bogs 3. In wells 4. In sites covered by rising seawater level V. Biomineralization (Phytoliths) 1. Opal 2. Calcium oxalate VI. Metal-oxide preservation 1. Near silver 2. Near copper or bronze 3. Near iron VII. Petrified remains 1. Siliceous mineralization 2. Calcareous mineralization

At fairly high temperatures (between 200 and 400 °C), carbonization causes characteristic deformations. In cereals, the most obvious changes are shrinkage in the length of the kernel together with a relative increase or ‘puffing’ in its circumference. Size reductions and specific patterns of swelling and/or crack-

11

ing appear also after the charring of certain seeds. Moreover, some organs do not generally survive charring (e.g. Boardman and Jones, 1989, Märkle and Rösch, 2008, Kislev and Rosenzweig, 1991). Thus, the seed coats in leguminous plants or the glumes and pales in cereals are only recovered on special occasions because they disintegrate into powder in most cases. The intensity of the deformation depends, among other things, on the amount of humidity present in the seed (the drier the grains, the less they are deformed), the spread of the heating, and the temperatures reached. Substantial information on the effects of heating on the seed of various plants has been gained experimentally by simulation of charring in laboratory ovens. Grains of various cereals and seeds of several pulses and flax have been the main elements tested. A determination of the amount of shrinkage in the seed of various crops also provides a better idea of the actual life-size dimensions of charred seed discovered in excavations (e.g. Märkle and Rösch, 2008, Braadbaart and van Bergen, 2005). Such experiments found that the degree of shrinkage or expansion of seeds vary according to the burning circumstances—temperature, time, degree of sealing from oxygen. Charred plant material is recovered from the excavated sediment either by direct collection or by separation techniques. There are lucky discoveries of hoards of burnt grains stored in containers or silos, which sometimes contain almost pure grains. In order to recover scattered remains embedded in site deposits, the excavator frequently resorts to separation by flotation. Water flotation is the simplest and cheapest technique, and usually separates the scattered charred remains present in the deposits effectively. This frequently includes relatively large amounts of cereal chaff and wood charcoal as well as other types of plant material that rarely appear in silos. The introduction of flotation in the late 1960s, and especially flotation machines, revolutionized archaeobotany by allowing excavators to search for seeds rather than rely on caches, and also improved the efficiency of such separation.

Impressions on pottery, daub, and bricks Imprints of grains and other plant parts on pottery contribute to documentation of crop plants in

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DOMESTICATION OF PLANTS IN THE OLD WORLD

archaeological sites. Such imprints are found particularly on handmade vessels. Pottery is one of the main diagnostic objects in archaeology, and imprints on pottery therefore have an obvious advantage, since once detected, they can be culturally classified and dated. However, imprints are frequently pressed into gritty, rough pottery (the common type of ceramics in early periods). On such a background the print is rather blurred, and unequivocal interpretation of such findings is often difficult. Daub and bricks provide another source of plant impressions. Straw, chaff, and similar dry plant material is often added to the wet clay to act as a tempering element. Plant parts can also become embedded in the clay by chance, and even if the organic matter does not survive well, the impressions remain intact in the dried or fired clay. They can serve as negative moulds for casting and reproducing the former inclusions.

effective preservatives, and plant remains in such places frequently retain their most delicate features. Excellent examples of waterlogged preservation have been found in lake-shore dwellings, submerged coastal areas, bottoms of old wells; as well as in the stomach contents of several human corpses retrieved from bogs in Denmark, Holland, and Germany. In some cases, the starch content of waterlogged-seed remains had vanished.

Preservation by oxides of metals Bronze, silver, and iron occasionally act as effective preservatives for plant material buried close to them. In humid situations they produce metal oxides, which impregnate the plant remains. Because copper-, silver-, and iron-oxides are highly toxic to bacteria and fungi, they block decomposition.

Mineralization Desiccated plant remains Preservation by desiccation, which blocks the processes of bacterial and fungal decomposition, occurs only under extreme dryness, so this source of evidence is confined to very arid areas. Such desiccated remains can be of particular importance because of their perfect preservation. Outstandingly rich remains of dried plants have been discovered in Egypt. There, grains, fruits, vegetables, corms, and other parts of plants placed in pyramids and tombs give an excellent account of plant cultivation in the Nile valley during predynastic and dynastic times. During the last decade several later sites, from the Iron Age onward, were discovered along the Red Sea. The finds include soft parts of vegetables, leaves, and flowers, which hardly ever survive under other conditions. Several discoveries of desiccated material were also made in caves in the Dead Sea basin.

Waterlogged preservation In Europe, valuable information has been obtained by examining plant material sunk in peat bogs or buried in the mud at the bottom of lakes, seas, or wells. Anaerobic conditions in these environments (and the presence of humic acids in bogs) act as

This type of preservation is brought about by filling of cell cavities by inorganic substances or by replacement of the content of cell walls by minerals. The most common is mineralization by calcium carbonate (CaCO3), silica, or phosphate. Seed coats and fruit shells of several plants undergo natural mineralization. For example, stones of hackberry (Celtis) contain large quantities of CaCO3 and the nutlets of several Boraginaceae accumulate silica. They sometimes survive in archaeological deposits without further means of outside preservation.

Phytoliths Plants deposit the mineral opal (SiO2.nH2O) between or within their cell walls, which creates minute silica bodies. As the silica is an inorganic material, it does not decompose in the soil/sediment and therefore, it is one of the most dominant botanical finds in archaeological excavations worldwide.

Digested or partly digested remains Preserved human feces (coprolites) constitute an only partly exploited source of evidence. Since

SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS

humans cannot digest cellulose, woody plant fragments, and shelled seed, they frequently retain their features after passing through the alimentary tract. Therefore when feces are charred, desiccated, or waterlogged, they often contain numerous identifiable plant fragments, which indicate the contents of the food in the tested human culture. Coprolite examination has already contributed significantly to American environmental archaeology. In the Old World this source of evidence has not yet been exploited extensively, although some results are already available (e.g. Hillman 1986; Dickson et al. 2000).

Chemical tests Tar compounds present in charred plant remains and organic residues precipitated in ancient vessels can be identified by gas liquid chromatography, infrared spectroscopy, and other tests used by organic chemists. Such detection is possible even when these substances survived in minute traces. Significantly, some of these chemical compounds are specific to a single crop species or a single plant product. They can be used as diagnostic traits for crop identification (e.g. Evershed 2008).

Evidence from the living plants Several principal contributions to the understanding of the crop-plant evolution are made by the study of the living plants. A major contribution is the identification of the wild progenitors from which the various domesticated plants could have been derived. Once the wild ancestry of the crop has been determined the following examinations can be carried out: (i) comparison of the cultivated varieties (‘cultivars’) with their wild counterparts in order to determine the main morphological, physiological, chemical, and genetic changes that took place under domestication; (ii) assessment of the range and the structure of genetic variation (chromosomal, protein, and DNA polymorphisms) present in the wild progenitor and those found in the domestic derivatives;

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(iii) assessment of changes in adaptation. Answers can be sought to the following questions: Which adaptations, that are vital under wild situations, have broken down under domestication? What are the new ‘syndromes’ that have evolved under domestication? Which selective forces are responsible for these changes? (iv) delimitation of the distribution areas of the wild progenitors. This often provides information on the place of origin of the crops. A second major contribution comes from examination of the crops and the ways they are handled, particularly under traditional (‘primitive’) systems of agriculture. Such studies include: (i) patterns of variation in each crop and their geographies; (ii) methods of cultivation and uses; (iii) genetic systems operating in the various crops and the breeding traditions used; and (iv) examination of the interconnections between the domesticated varieties and their wild relatives. The role of weedy races accompanying the crops is significant for this topic. The following sections survey the main tools used for identification of wild progenitors. They also deal with some of the complications and problems involving the use of the wild relatives for elucidation of crop-plant evolution.

Discovery of wild progenitors A principal goal in the study of the living plants is the identification of the wild progenitors of the crops; that is, the wild stocks from which the domesticated plants could have evolved. Plant domestication is a relatively recent evolutionary event. Therefore, one can expect that most wild ancestors are still alive and include forms similar to those that existed in pre-agricultural times. Indeed, the wild progenitors of the majority of the world’s main food plants have already been identified. Many of them became known only during the last thirty-five years. Several complementary tests are available for the identification of the wild progenitors of crops. They

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DOMESTICATION OF PLANTS IN THE OLD WORLD

all seek to determine which of the wild species, usually grouped together with the crop in the same biological genus, is most closely related to the domesticated plant. (i) The classical taxonomic approach recognizes the wild progenitor by its close morphological resemblance to the crop. It sorts out the wild progenitor from among all the other wild taxa grouped in the crop’s genus by its closest morphological and anatomical affinities to the cultigen. This is the oldest method. In some cases morphological comparison provides sound clues for the determination of ancestry. However, in numerous cases the wild background is taxonomically complex. Moreover, many crops exhibit a bewildering morphological variation, very different from the patterns present in wild plants, and this can be confusing in relationship analysis. Critical evaluation in such cases necessitates genetic verification; which can be obtained through cytogenetic analysis and by comparative molecular (DNA and/or protein) tests. (ii) Cytogenetic analysis aims at elucidating the chromosomal affinities between the domesticated plant and the wild species. It also tests whether or not these wild taxa are separated from the crop (and isolated from one another) by hybrid sterility or other reproductive isolation barriers. Since evolutionary domestication is a recent development, the crop and its wild progenitor should retain a considerable amount of homology in their chromosomes. In contrast, other species grouped in the genus were probably formed long before the beginning of agriculture. As a result, they could have diverged considerably in their chromosomal constitution. The principal tool of cytogenetics is a program of crosses between species followed by examination of inter-specific hybrids. Chromosome pairing in meiosis indicates the degree of chromosomal homology between the two parents. In most crops (particularly in grain crops), the cultivars show full homology and complete inter-fertility with only one of the wild species in the tested genus. Such a wild (congeneric) type is recognized as the ancestor (wild progenitor) of the crop. Together they comprise the ‘primary’ gene pool of the crop. In contrast, other members of the genus are frequently chromosoma-

lly distinct and are separated from the crop by strong reproduction isolation barriers such as crossincompatibility, hybrid inviability, or hybrid sterility. Such species are often called ‘alien species’ and their chromosomes ‘alien chromosomes’. They comprise the ‘secondary’ and ‘tertiary’ gene pools of the crops (Harlan and de Wet 1971, p. 107; Harlan 1992a). To summarize, fully fertile hybrids showing normal chromosome pairing in meiosis point to close genetic relationship between the tested parents and implicate the wild plant in the ancestry of the crop. Lack of chromosome homology and the presence of strong reproductive isolation barriers indicate longestablished genetic divergence and rule out the tested wild plant from being a progenitor of the crop. Chromosome analysis of domesticated plants has frequently also to deal with complications due to polyploidy; i.e. the formation of new subspecies (or even new species) by doubling of chromosome numbers. Evolution by polyploidy is common in the plant kingdom. Many wild plants (including progenitors of domesticated plants) are not standard diploids but polyploid entities. One class comprises auto-polyploids which increased their chromosome number from the standard of two dosages (diploid condition) to three sets (triploids), four sets (tetraploids), or even higher levels. Such increases are not uncommon among vegetatively propagated crops (corm and tuber plants, ornamentals, and some fruit trees). A second class includes allo-polyploids; i.e. types formed by inter-specific hybridization followed by chromosome doubling. This combines the genetic contents of two (or even more) donor species in a new hybrid species. Bread wheat is a product of such fusion under domestication (pp. 47–48). Cultivated tobacco and the New World cottons had a similar mode of origin. In such crops, a special cytogenetic test known as ‘genome analysis’ helps to elucidate the polyploid origin and to identify the parental stocks, which donated their chromosomes to the new polyploid entities. (iii) Advances in molecular biology provide critical tools for assessing the range of variation, and finding the structuring of genetic variation (genetic polymorphism) in crops and their wild relatives. This information can be used for determining the genetic

SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS

affinities and phylogenetic relationships between the cultivars and their wild relatives. As major breakthroughs in this field occurred since the publication of the third edition of this book some ten years ago, it receives extensive treatment in the present edition. Since the 1960s, critical results have been obtained by testing protein variation in crops and their wild relatives, particularly enzyme variants (isozyme and allozyme polymorphism) and/or variability in storage proteins deposited in seeds (Soltis and Soltis 1989). Proteins are the primary products of the genes, and therefore their variability reflects differences in the hereditary material. Gel electrophoresis separation makes it possible to discern variation and differences in numerous proteins. Lately, detecting protein variability has become an outdated technique although it is time- and cost-efficient. Its relatively low output makes this technique less attractive than DNA-based marker systems. Since the late seventies, large-scale analysis of DNA variation became possible in a wide range of techniques. The first major breakthrough for variability analysis was the development of restriction fragment length polymorphism (RFLP) technology. Restriction enzymes cleave the DNA strands at specific sites (‘restriction’ sites’) into identifiable fragments. Individuals with identical site arrangements yield identical DNA fragments, while those that carry mutations in these sites produce different fragment patterns. Because restriction sites along the DNA strands of genes are numerous, polymorphism in these sites is enormous both within and between populations of crops and their wild progenitors (e.g. Havey and Muehlbauer 1989). Another major breakthrough was made by development of the polymerase chain reaction technology (PCR), by which quantities of DNA fragments, large enough for variation analysis, are amplified from minute samples of target DNA and their specific primers. Indeed, PCR is used in almost all DNA polymorphism analyses. In the mid-nineties, combining PCR and RFLP resulted in a most popular technique, namely AFLP (amplified fragment length polymorphism, Mueller and Wolfenbarger 1999). Although it is still relatively expensive, this method is highly reliable and informative, and indeed many of the recent elucidations on crop origins such as

15

wheats, barley, and chickpea were available through this procedure (Heun et al. 1997b; Badr et al. 2000; Özkan et al. 2002; Nguyen et al. 2004; Duc et al. 2010). Another technique that uses arbitrary primers for the PCR and needs minute amounts of genetic material is random amplified polymorphic DNA (RAPD). It is a straightforward method, but is relatively less reliable and informative. However, it reveals genetic diversity easily, as found, for example, in flax and in lentils (Sharma et al. 1996; Fu et al. 2002). Another highly informative technique known as microsatellites, involves short DNA repeats that show high variation in repeat-number between individuals (Varshney et al., 2005). Also known as SSR (simple sequence repeat), this method produces highly unique patterns that in fact are the basis for individual DNA fingerprinting in many different organisms including crops (Molina-Cano et al. 2005) and humans. Direct DNA sequencing of the genetic text at target sites may also reveal variability between individuals. In general, it retrieves phylogenetic information from diversity in specific loci in the DNA, as manifested (for example) in the sad2 locus in flax (Allaby et al. 2005), or the btr1/2 loci in barley (Komatsuda et al. 2007). Remarkable advances in sequencing technology have recently enabled comparisons of large DNA sequences that accordingly embed variations or point mutations in the form of single nucleotide polymorphisms (SNPs, Rafalski 2002). Such point mutations are abundant throughout the genome. They reflect DNA variability and were used recently to study diversity between and within crops and their ancestors as done for example in barley (Kanazin et al. 2002). Today, the bewildering array of DNA markers and the ability of automatic sequencing of genes of interest in hundreds of individuals can be carried out in a very short time. This revolutionized our ability to trace and assess genetic and evolutionary relationships and construct reliable phylogenetic trees for crops and their progenitors.

Distribution of the wild progenitors The wild relatives can frequently provide critical information about where domestication occurred.

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DOMESTICATION OF PLANTS IN THE OLD WORLD

In many crops, the progenitors occupy limited geographic territories—much smaller distribution areas than those of their domesticates. Because domestication is a recent development, it is safe to assume that the distributions of the wild forms (weeds excluded) have not undergone drastic changes since the beginning of cultivation. Delimitation of the wild relative’s distribution thus marks the territory in which the crop could have been taken into cultivation. The narrower the distribution area, the more accurate the placement. Fortunately, the distribution of the wild progenitor of emmer wheat—a principal Old World ‘founder crop’—is confined to the Fertile Crescent (Map 4, p. 42). It is thus possible to plot, fairly accurately, the area where Neolithic agriculture could have started. The archaeological records have fully corroborated this supposition. The delimitation of the place of origin of the chickpea is even more precise. Its wild ancestor is endemic to south-east Turkey (Map 10, p. 88). However, not all wild progenitors have such a limited distribution. Some (e.g. the wild relatives of the foxtail millet, oat, flax, and numerous fruit trees) are distributed over extensive territories. The use of their distributions for the determination of places of origin is much less accurate.

Weeds and domestication Some crops seem to have entered cultivation not directly but by first evolving weedy forms. The establishment of tilled fields (as well as other disturbed habitats) gave an opportunity to numerous unwanted plants to invade the newly made habitats and to evolve as weeds. Weed evolution went hand in hand with crop cultivation and from the very start the control of these invaders seems to have been a major problem in agriculture. Noxious weeds are plants that have successfully adapted themselves to the ecology of the tilled ground. They are independent only because they retain their wild mode of seed dispersal, and germinate and develop in spite of the efforts of the cultivator to eradicate them. But if any such weeds turns out to produce a valuable commodity, it can eventually change its relationship with humans. The cultivator may follow the rule ‘if you can’t beat them, join them’, and start to utilize the weed by intentionally planting its seed, harvesting its fruits, and selecting the better

yielders. Several Old World crops are such ‘secondary crops’, i.e. plants that entered domestication through the back door of weed evolution (Vavilov 1949–50, 1987). They were added to the crop assemblage only after the establishment of the principal seed crops. Well-documented cases are those of the oat, Avena sativa (pp. 66–69), and of the gold of pleasure, Camelina sativa (p. 111). Several other plants seem to have followed a similar evolution under domestication.

Classification and botanical names Orientation in crop plant evolution is frequently complicated by inconsistencies in species delimitation and by proliferation of botanical names. As already noted, cultivated plants are, as a rule, very variable. Furthermore, evolution under domestication commonly involves drastic modifications in organs and traits that stay fairly uniform in wild plants. Traditional taxonomic treatments of crops suffered from over-splitting, since they were based almost entirely on morphological comparisons. Frequently, inter-fertile crop varieties were ranked as separate species and called by different botanical names because they looked so different. For example, classical cereal taxonomists recognized twelve to fifteen species of cultivated wheats (see Table 3, p. 29). Barley and common oat were each split into two or more species (Table 5, p. 57). Similar splitting and species ranking characterized numerous other crops. With the accumulation of cytogenetic information, it has become increasingly clear that the traditional classification of many crops is inadequate and even misleading. Frequently two, three, or even half a dozen ‘species’ were found to be inter-fertile, chromosomally homologous, and genetically interconnected. Moreover, in many cases the conspicuous morphological distinctions turned out to be governed by single mutations (Table 7, p. 61). Ranking such types as independent species is unjustified. They represent only varieties within species and deserve only intra-specific ranking. In wheats, modern taxonomic revision has reduced the species number to five (Table 3, p. 29). All cultivated barleys are grouped in a single species (Table 5, p. 57), as are all common oats. The discovery of the wild progenitors necessitated another nomenclature change. Because the

SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS

wild plants and their cultivated derivatives are genetically interconnected, they cannot be regarded as fully diverged species. According to internationally agreed taxonomic rules, once a wild ancestor is satisfactorily recognized, the crop and its wild relative cease to be treated as two separate species. Instead, they should be lumped in a single collective species, frequently also including related weed types. In other words, the wild and crop types are considered as subspecies or varieties of a single biological species and botanically named accordingly. However, habits die hard. Old names and traditional classifications are still widely used by many researchers. Wild progenitors, in particular, are commonly referred to as independent species. To avoid confusion, botanical orientation in crops should begin with the following questions: (i) What are the main cultivated, weedy, and wild elements in the crop complex? (ii) What botanical names are used by different people for these intra-specific taxa of the crop complex? (iii) What are the other fully divergent (‘alien’) species placed in the same genus?

Radiocarbon dating and dendrochronology Radiocarbon (14C) dating was developed by W.F. Libby at the University of Chicago soon after the Second World War and created a real breakthrough in archaeology (Libby et al. 1949). Previously, one could date archaeological remains only by relative chronology based on stratigraphy and cultural associations. The introduction of radiocarbondating methods brought about absolute dates and made possible age comparisons between cultures in the various parts of the world. Until the 1980s, radiocarbon dating demanded relatively large samples of charred material. Most tests were made on carbon sources (such as wood charcoal or hoards of grain) obtained from secured archaeological contexts. By then, radiocarbon accelerator mass spectrometry (AMS) technique had been developed (Nelson et al. 1977). This technique, based on counting the atoms instead of the decay product, made it possible to reduce the size of the sample to a few milligrams.

17

Consequently, it became possible to date accurately individual charred grains or to reduce the amount of material removed from rare or very important objects. The application of the AMS test is sometimes critical (Harris 1986). Beside dating, radiocarbon serves to detect—and eliminate—errors in chronologically associated samples to the wrong strata or archaeological level. These errors are due to intrusion; i.e. the occasional displacement of plant remains from one layer or context to another as a result of boring by animals or other interferences (Boaretto 2007, 2009). Better precision and accuracy of archaeological dating brought about by several improvements in this field: (i) the improved techniques of chemical pre-treatment for the elimination of contaminations from the sample material; (ii) the ability to reduce the size of the sample to few milligrams, extend the range of possible material to be dated, the care taken to date archaeological finds from secure, sealed, contexts; (iii) the building of a calibration curve up to the range of radiocarbon dating’ 50,000 year (Reimer et al. 2009). The principle of radiocarbon dating is based on the measurement of the radioactive 14C isotope concentration in sample material, in comparison to the stable carbon isotope 12C. The production of 14C, due to the interaction of cosmic rays with molecules, takes place in the atmosphere as neutron absorption on 14N atoms. These 14C forms CO2 molecules, are then introduced into different natural reservoirs (e.g. hydrosphere, biosphere) by different physical and chemical processes. In these reservoirs the relative 14C and 12C concentration is in equilibrium as long as the exchange with the atmosphere stays open. When this exchange ceases (e.g. when the organism dies), the 14C concentration starts to decrease due to decay of the radioactive 14C. Therefore, the measured concentration of 14C in material examined depends upon the time elapsed from when the exchange with the atmosphere stopped. The radiocarbon age is calculated from the measured 14C concentration using the decay law. The smaller the proportion of 14C in the tested organic remains, the older the sample.

18

DOMESTICATION OF PLANTS IN THE OLD WORLD

The concentration of 14C in the atmosphere was not constant in the past (Suess 1970), and some fluctuations seem to have occurred in the concentration of this isotope in the atmosphere. This means that age estimates based on conventional radiocarbon timescale are in need of some calibration. More precise dating was made possible by establishing the sequences of annual rings in wood remains of trees (oaks, bristlecone pine), and currently also in corals; and radiocarbon dating of the rings in these sequences. Recently, these sequences have been updated and extended to 50,000 years BP (Reimer et al. 2004). By plotting radiocarbon ages against treering ages, calibration curves have been constructed correlating radiocarbon dates with dendro- or calendar times (Fig. 1). Thus by means of dendrochronology, the radiocarbon timescale can be calibrated against annual tree-ring chronology to calendar dates (Stuiver et al. 1986; Stuiver and Reimer 1993; Baillie 1995). Joined by other dating methods (U-Th ages from corals), calibration reaches

the limit of radiocarbon dating which is 50,000 years (Reimer et al. 2009). A relevant part of the current calibration curve is shown in Fig. 1. One can see that except for the last three thousand years, the radiocarbon age represents somewhat reduced estimates. Moreover, the differences increase in time. Thus for radiocarbon dates 3,000–4,000 years before present (BP), calibration adds 200–400 years. For older radiocarbon dates (6,000–9,000 BP), the addition is already 700–1,000 years. For still older radiocarbon dates the calibration differences are even greater. The result of the calibration process is usually presented in the form of probability distribution of the radiocarbon age (black curve) as it is shown in Fig. 2. The radiocarbon determination, with one standard deviation, is given in the center of the upper side of the plot. The distribution of the radiocarbon determination (red curve on the left-hand Y axis) is projected on the calibration curve (an uneven double blue line on the X axis, in calibrated years BP), to produce the radiocarbon age (solid

Atmospheric data from Reimer et al (2004); Oxcal v3.10 Bronk Ramsey (2005); cub r:5 sd:12 prob usp[chron]

Radiocarbon determination

25000BP

20000BP

15000BP

10000BP

5000BP

0BP 25000CaIBP

20000CaIBP

15000CaIBP

10000CaIBP

5000CaIBP

Calibrated date Fig. 1 The radiocarbon calibration curve for the past 25,000 years based on trees and corral annual ring sequences. Adapted from OxCal 3.10 software Bronk-Ramsey 2005 (Bronk-Ramsey 1995; Bronk-Ramsey 2001). The straight thin line represents the ideal 1:1 correspondence between radiocarbon age and calendar age assuming constancy of 14C concentration in the atmosphere. If 14C years were the equivalent to calendar years, all of the data would fall on the diagonal straight line.

SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS

19

Atmospheric data from Reimer et al (2004); Oxcal v3.10 Bronk Ramsey (2005); cub r:5 sd:12 prob usp[chron]

9200±45BP

9500BP

68.2% probability 10420BP (68.2%) 10260BP 95.4% probability 10500BP (95.4%) 10240BP

Radiocarbon determination

9400BP 9300BP 9200BP 9100BP 9000BP 8900BP

10800CalBP

10600CalBP

10400CalBP

10200CalBP

10000CalBP

Calibrated date Fig. 2 Presentation of radiocarbon results—probability distribution of a sample that was measured 9200±45 14C year BP (=Before Present); see text for explanations of this result. The calibration is performed using the software OxCal 3.10 Bronk-Ramsey 2005 (Bronk-Ramsey 1995; Bronk-Ramsey 2001).

black distribution). The probability of dates is given in the top-right corner in two different levels of confidence, 68.2% probability (±1σ) and 95.4% probability (±2σ). For each probability, the range of dates is given, from left to right, with the percentage in the middle representing how much this range covers. When there is more than one range of probabilities for each 1 or 2σ probabilities, each interval appears separately. A straight line is placed underneath the black curve to make it easy to determine the dates as can be seen on the X axis. In case of more than one range of probabilities for each 1 or 2σ probabilities, such straight line is placed under each curve. It is important to note that radiocarbon dates are given as a probability range, which means the actual dates could be between two possible dates, rather than the median dates between them. In addition, as Figs 1 and 2 show, calibration curve is far from being 1:1 straight line, and several wiggles and flat areas can be seen. As a result, due to the fact they fall into a flat area in the calibration curve, several different radiocarbon dates can produce similar

ranges. Disturbing such area occur around 10,000 uncal BP—a period of great importance for the understanding of the beginnings of agriculture. These areas in the calibration curve are still widely misunderstood aspects of radiocarbon dating. As a rule, dates mentioned in this book are calibrated radiocarbon dates before present (cal BP). Therefore, the use of ‘BP’ and ‘cal BP’ denote calibrated 14C dates (years before present, i.e. before 1950). BC (years Before Common Era/Before Christ) denotes uncalibrated dates, either from Bronze Age and later radiocarbon dates, when calibration has little impact, when dates drawn from traditional methods of stratigraphic dating, or when calibrated dates were unavailable. Also, progress of evidences in this book is presented on a timetable moving from older to newer dates, from earliest to recent finds. Therefore, when we separate millennia to first and second halves (i.e. ‘the second half of the eleventh millennium cal. BP’ or ‘the first half of the ninth millennium cal. BP’), the first half will be older and the second one,the younger.

C H A PTER 3

Cereals

Cereals (annual grasses grown for their grains) have been the principal crops of most Old World’s civilizations. Their grains constitute the main source of calories for mankind. Different parts of the world depended on cultivation of different cereals as staple foods. Food production in the Mediterranean basin, Europe, the non-tropical parts of Asia, and (to some extent) the highlands of Ethiopia, was based primarily on wheat and barley. South and south-east Asia had rice, broomcorn, and foxtail millets. America grew maize. Africa south of the Sahara raised sorghum, pearl millet, and several other native grasses. Cereals thrive on open ground (the cultivated field) and usually complete their life cycle in less than a year (Plate 1). The nutritive value of their grains is generally high, and the seed can be stored for relatively long periods. In most cereals the grains are packed with starch, and in some, such as in wheats or oats, they also contain an appreciable amount of protein. Compared with grain crops belonging to other plant families (e.g. legumes), yields in cereals are relatively high. However, they depend heavily on soil fertility and the availability of fixed nitrogen. Wheats and barley are the principal ‘founder crops’ that started food production in south-west Asia and Europe. The first definite signs of wheat and barley domestication appeared in the Fertile Crescent in the second half of the eleventh millennium cal BP. Later, grains of these cereals constituted the bulk of plant remains retrieved from south-west Asian Neolithic, Chalcolithic, and Bronze Age contexts. Wheats and barley were the main domesticates that led the explosive expansion of the Neolithic 20

agriculture from its ‘core area’ to the vast territories of west Asia, Europe, and north Africa. Several other grasses entered agriculture in west Asia and Europe, but apparently, only after the firm establishment of the ‘first wave’ in south-west Asia. Such an early domesticate is broomcorn millet, followed by common oat. Two major ‘alien’ cereal crops, namely rice (native to south-east Asia) and sorghum (native to Africa south of the Sahara) arrived as well, sometime in Hellenistic or Roman times. Apparently, their establishment in Mediterranean agriculture was much later. Wheats, barley, and several other prominent cereals, differ from the majority of flowering plants in their system of pollination. While the great majority of the world’s plant species are allogamous (crosspollinated), most domesticated seed crops are predominantly self-pollinated, or ‘selfing’ (Zohary 1999). Significantly, selfing is also a characteristic feature of their wild progenitors. In wheat, barley, and most other grasses, it is brought about by precocious shedding of the pollen inside the florets; that is, before they open. Was it a mere chance that the first grain plants that were successfully domesticated in the Old World were selfers? Several facts suggest that predominantly self-pollinated plants were better suited to domestication than cross-pollinated candidates. One major advantage of self- over cross-pollination in incipient domesticants is the fact that selfing automatically brings about reproductive isolation between the crop and its wild progenitor. This isolation barrier enables the farmer to grow a desirable cultivar in the same area in which its wild relatives abound, without endangering the cultivar by

CEREALS

genetic swamping. If indeed domestication of south-west Asian cereals started in areas where the wild progenitors were common, genetic separation between tame and wild could have been a major asset. Under cross-pollination the initial small patches of cultivation would have been exposed to large quantities of pollen grains from its wild relatives. Safeguarding the identity of the domesticated varieties under a cross-pollination system would have become difficult or even impossible (Harlan 1995b). A second advantage of self-pollination lies in the genetic structure maintained within the crop. Selfing results in splitting the crop’s gene pool into inbred homozygous lines. Variation is thus structured in the form of numerous true breeding cultivars. Since these cultivars are automatically ‘fixed’ by the pollination system, they can easily be maintained by the farmer, even when planted side by side. In contrast, the preservation of varietal identity in crosspollinated plants is much more problematic. It requires repeated selections towards the desired norms, constant care to avoid the mixing of types, and the prevention of pollination from undesirable plants. It is therefore not surprising that early successes in plant domestication involve selfers. In fact, the ‘first wave’ domesticants in south-west Asia, namely emmer wheat, einkorn wheat, barley, pea, lentil, chickpea, bitter vetch, and flax, are all selfpollinators. Cross-pollinated crops entered agriculture later and comprise only a small minority among the traditional grain crops. Rye and faba bean are the only important cross-pollinated grain crops indigenous to south-west Asia. It is noteworthy that wheats, barley, and all other Fertile Crescent ‘founder crops’, are not obligatory selfers but predominantly self-pollinated plants in which rare events of cross-pollination occur. Such pollination systems are admirably suited for rapid grain-crop evolution, because, being annual, they produce new types recurrently. Occasional crosspollination provides the crop with genetic flexibility and serves to combine and reshuffle genes from different sources. The numerous cycles of inbreeding that follow the rare events of crossing lead to the fixation of many new recombinant lines. The grower can easily pick up the more attractive ones among them.

21

Cereal cultivation comprises several essential stages: sowing seeds in tilled fields; weeding; reaping the mature spikes or panicles; threshing; winnowing; and storage. The introduction of these practices automatically initiated drastic changes in grasses taken into cultivation, setting them apart from their wild progenitors (Harlan et al. 1973; de Wet 1975). Most conspicuous is the selection for types in which the mature dispersal units are retained on the mother plant, and in which the wild-type adaptation for seed dissemination has broken down (Zohary 1969). In cereals, this implies a shift from shattering ears or panicles (in wild forms), to non-shattering types (in domesticated forms). In the wild forms, survival depends on seed dispersal and the various wild cereals have evolved elaborate seed dissemination devices. In wild wheats (pp. 34, 40) and wild barley (p. 53), the seed-dispersal unit (diaspore) comprises a single internode of the ear. Disarticulation of the spike’s rachis is thus an essential element of the wild-type seed dispersal. Plants in wild populations are constantly selected for quick shattering of their mature ears. In contrast, the introduction of planting and harvesting brings about automatic selection in exactly the opposite direction. Under the new system, a sizeable proportion of the seed produced by plants with brittle ears will shatter and would not be included in the harvest. In contrast, practically all grains produced by non-shattering mutants ‘wait on the stalks’ to be reaped by the grower. Therefore, under cultivation, non-shattering individuals have a much better chance to contribute their seeds to the subsequent generation. Consequently, non-brittle (or less brittle) mutants (which were disadvantaged under wild conditions) become highly successful under the new human-controlled system. Thus, when wild cereals are brought into cultivation, one should expect selection for non-brittle forms whether or not the cultivator is aware of this trait. Furthermore, the incorporation of non-brittle mutants makes the crop fully dependent on humans, as non-shattering plants lose their seed-dispersal ability, and can no longer survive under wild conditions. Thus, a symbiotic relationship has been established between humans and their crops (Rindos 1984).

22

DOMESTICATION OF PLANTS IN THE OLD WORLD

The appearance of non-shattering mutants in cereal crops was probably a swift process. Genetic considerations indicate that such a shift could have been accomplished in the course of a few dozen generations of selection (Zohary 1969; Hillman and Davies 1999; Kislev 2002; Tanno and Willcox 2006a; Fuller 2007). It is our view, however, that much more research is needed in order to increase our understanding of the shifts between wild and domesticated populations, and the speed of their development. To date, the most effective (and easy to trace) diagnostic indication for domestication in archaeological cereal remains is the rough disarticulation scars in the internodes (Plates 2 and 3). This shift is controlled mostly by a recessive mutation in a single gene or by two such mutations (Table 7). As Kislev (1989b) noted, some lower-ear spikelets of wild cereals, up to 10% of them, do not disarticulate naturally. This factor had to be taken into account while discussing the domesticated status of a given archaeobotanical assemblage. Tanno and Willcox (in press) re-examined recently the spikelet bases from eleven Turkish-Syrian sites dated to the PPNA-PPNB period. They draw attention to the difference between upper and lower scars, between wheat and barley, and especially noted that only when the abscission scar is well preserved it can be use as a diagnostic tool. A less reliable diagnostic trait is grain size. Seeds in cultivars are usually plumper and larger than those produced by their wild relatives. However, the increase of grain size evolves slowly from domestication onward. As Nesbit (2002) noted, grain’s shape and size differences are clearly visible in Pottery Neolithic find, but less clear in the PrePottery Neolithic. Recently, Gegas et al. (2010) explored the genetic basis of the variation in wheat grain size and shape. Their work demonstrates that ‘grain size has progressively increased through alterations both in grain width and length, followed at later stages by modifications in grain shape’. Therefore, size increase in grains (under cultivation) is not a fully reliable indication of the early stage of domestication. A third major outcome of introducing wild cereals into the regime of cultivation is the breakdown of the wild mode of seed germination. This

breakdown is manifested in the reduction of germination inhibition from dormant to synchronous germination. Most wild grain crops, especially annuals growing in Mediterranean-type (dry summer, wet winter) or in semi-desert climates, depend for their survival on the regulation of germination. A common adaptation is a delay of germination from the time of seed maturation until the onset of favourable conditions in the next growing season. In the Mediterranean basin this means germination inhibition—even if the seed is wetted—from the time of seed maturation in early summer to the start of the rainy season in the following autumn. Another adaptation is the delay of germination of grains from the same dispersal unit over two or more years. This protects the populations from the crippling effects of dry or otherwise bad years. In wild emmer wheat the dispersal unit contains two kernels. One germinates in the ensuing autumn; while in the second kernel, germination is frequently delayed until the following year. Wild barley diaspores contain each only a single seed, but not all dispersal units germinate in the first winter. Under cultivation, this wild-type regulation of germination is no longer advantageous for the farmer, and mutants in which germination inhibition has broken down are selected automatically. This type of selection has evidently been operative in the Old World domesticated cereals. Most cultivars have lost the germination-inhibition patterns which characterized their wild progenitors, so that all their seeds germinate at the same time whenever planted by the farmer. Several other traits that characterize domesticated cereals (the ‘domestication syndrome’) seem to be the outcome of conscious and/or unconscious selection under domestication (for review see Zohary 1969; Harlan et al. 1973). They include: (i) selection towards erect plants, synchronous tillering, and uniform ripening; (ii) increase of seed production by addition of fertile florets and/or increase in the size of the inflorescence or the number of ears or panicles produced per individual plant; (iii) decrease of awns, of glumes’ thickness, and investment of grains (from hulled to naked grains).

CEREALS

Wheats: Triticum Wheats are the universal cereals of Old World Mediterranean-type agriculture. Together with barley, they constitute the principal grain stocks that founded Neolithic agriculture, and the main element responsible for its successful spread. Since this early start, wheats have retained their crucial role in Old World food production and have given rise to numerous advanced forms. As of 2008, wheats and rice rank second in the world’s grain production, whereas maize rank first (http://faostat.fao.org/ site/339/default.aspx). Wheats are grown extensively throughout the temperate, Mediterranean, and subtropical parts of both hemispheres.1 Wheats are superior to most other cereals (e.g. maize, rice, sorghum, or barley) in their nutritive value. Their grains contain not only starch (60–80% carbohydrate), but also significant amounts of protein (8–14%). The gluten proteins present in the grain give wheat dough its stickiness, and its ability to rise when leavened. In other words, the gluten proteins bestow the dough with unique rising and baking qualities. Wheats were, and still are, the preferred staple food of traditional farming communities throughout the Old World. Thus, it is not surprising that in numerous cultures, the idea of food has been equated with bread. Several distinct wild species of the genus Triticum L. were introduced into domestication. Most modern wheat cultivars belong to two biological species: (i) bread wheat, Triticum aestivum (2n = 6x = 42 chromosomes), valued for the baking of high rising bread; and (ii) hard or durum-type wheat, T. turgidum, (2n = 4x = 28 chromosomes), used for the preparation of pasta and low-rising bread. Other Triticum species, as well as more primitive cultivars of the above-mentioned two species, were important in the past and survive today only as relic crops. All wheats are almost fully self-pollinated. Therefore, variation in these cereals is moulded in the form of numerous inbred lines.

23

Chromosomally, wheats comprise a polyploid series (for review of wheat cytogenetics, see Riley 1965; Sears 1969; Miller 1987; Feldman et al. 1995). Some domesticated forms have a diploid chromosome number (2n = 2x = 14 chromosomes) and contain two sets of a single genome (designated AA). Other wheats are tetraploid (2n = 4x = 28) and combine two distinct genomes (either BBAA or GGAA). Others, are hexaploid (2n = 6x = 42) and contain three different genomes (BBAADD). Domestic wheats fall into five chromosomal groups: two diploid wheats, two tetraploid wheats, and a single hexaploid wheat. Forms within each group are interfertile and share the same chromosome constitution. In contrast, hybrids between groups are highly sterile. The taxonomic classification of domesticated wheats and their closely related wild types (Van Slageren 1994) is based on these cytogenetic criteria. The following five biological species (Table 3) are recognized today in the genus Triticum: 1. Diploid T. monococcum L. or einkorn wheat (genomic designation AmAm) comprises both wild and domesticated forms (Fig. 3). Domesticated einkorn with its characteristic hulled grains was an important grain crop in the past. It survives today only as a relic crop. 2. Diploid T. urartu Tuman. ex Gandiljan. (genomic designation AA). This diploid wheat comprises only wild forms and is largely confined to the Fertile Crescent belt and extends to Armenia (Waines 1996; Valkoun et al. 1998). Apparently, it contributed its diploid set of chromosomes (long before domestication) to the 4x hard wheat, T. turgidum, and the 6x bread wheat, T. aestivum (Dvořák et al. 1998; Dvořák et al. 1993, 1988). Morphologically, T. urartu closely resembles the wild forms of T. monococcum. However, crossing experiments indicate that T. urartu is reproductively isolated from both wild and domesticated einkorn wheats. Interspecific F1 hybrids between these wheats are malesterile and do not set seeds upon self-pollination

1 Excluding the closely related Aegilops L., which comprises some twenty-two species. Some Aegilops species show close cytogenetic links with the wheats. Indeed, diploid Aegilops species participated in the formation of the polyploid Triticum species. For this reason several researchers (see, for example, Sears 1969; Morris and Sears 1967) lump together both genera, and include the Aegilops species within Triticum. Others (e.g. van Slageren 1994; Zohary 1965) keep the traditional classification, but consider Triticum and Aegilops as a single natural group (the ‘wheat group’).

24

DOMESTICATION OF PLANTS IN THE OLD WORLD

(Waines and Barnhart 1992; Waines 1996; Valkoun et al. 1998). Also isozyme and DNA tests (Jaaska 1997b) indicate that these two diploid wheats are distinct. 3. Tetraploid T. turgidum L. (genomic designation BBAA) comprises (see Table 3) wild emmer wheat, domesticated emmer wheat, durum wheat, and several other domesticated tetraploid forms (Figs 4 and 5, and Plates 2–4). Cytogenetic and molecular tests have shown that the genome A of tetraploid T. turgidum is closely related to the diploid genome present in T. urartu (Dvořák et al. 1993, 1988; Jaaska 1997b). In contrast, the genome of diploid T. monococcum is more remote. Consequently, the classical explanation of the polyploid origin of tetraploid T. turgidum had to be revised. Now it is clear that a T. urartu-like diploid wheat was involved in the polyploid origin (in the wild) of tetraploid T. turgidum subsp. dicoccoides; while diploid wild T. monococcum—long considered as the donor of the A genome to wild emmer wheat—had little to do with its polyploid origin. From the start of agriculture, hulled (= glumed) emmer wheat emerges as the principal stock of domesticated wheat. It gave rise to the wide range of present-day, free-threshing, tetraploid durum-type wheats. By hybridization with a wild Aegilops species, domesticated tetraploid T. turgidum formed an additional species: the hexaploid bread wheat (see below). 4. Tetraploid T. timopheevi Zhuk. (genomic designation GGAA) includes both wild and domesticated hulled forms. Domesticated Timopheev’s wheat is endemic to a small area in Georgia, and seems to represent only a local episode in wheatcrop evolution. The A genome in T. timopheevi is most similar to the genome present in diploid T. urartu indicating that an urartu-like diploid ancestor was probably involved in the polyploid formation (in the wild) of timopheevi’s wheat. 5. Hexaploid T. aestivum L. or bread wheat (genomic designation BBAADD) originated under cultivation by the addition of the DD chromosome complement of the wild grass Aegilops tauschii Coss. [= Ae. squarrosa L.] to the domesticated tetraploid BBAA turgidum wheats. The extraordinarily variable T. aestivum group constitutes the most important wheat crop of today (Fig. 6). It comprises several primitive, hulled, spelttype wheats, and numerous free-threshing forms (including modern bread wheat).

Traditional wheat classification has suffered from excessive splitting. Formally, wheat taxonomists regarded every main morphological type in the domesticated wheats as an independent species and recognized more than a dozen distinct species in the genus Triticum (for details, see the treatments of Percival 1921; Schiemann 1948; Dorofeev et al. 1979). However, in view of the accumulated information on genetic affinities between types, such species delimitation is no longer justified. Today, wheats are usually grouped in five biological species. Table 3 (p. 29) lists the various traditional species names in Triticum together with the modern grouping of the wheats. Table 4 presents the wild stocks from which the principal domesticated cereals were derived and summarizes the main evolutionary events that led to the formation of the crops. Domesticated wheats fall into several distinct classes according to their response to threshing: 1. The more primitive forms, i.e. diploid einkorn, tetraploid emmer, and hexaploid spelt have hulled grains. Their grains are enclosed in the spikelet by toughened glumes that do not break during threshing. In hulled wheats, the products of threshing are spikelets, not grains (Fig. 7). More advanced domesticated wheats, i.e. tetraploid durum-type and hexaploid bread wheats, are free-threshing. Their glumes are thinner and do not invest the grains tightly. Threshing releases the naked kernels (e.g. Fig. 5, 6). Because of this difference in threshing product, the handling of hulled wheats by the farmer is different from that of the free-threshing ones. In the first, the spikelets are stored and marketed as such, and the grains have to be freed (usually by pounding) before they can be used. The utilization of naked wheats is simpler. After threshing, the free grains are winnowed, sieved, and then stored ready for milling. Because of the different appearance of the marketed products, hulled and free-threshing wheats were often regarded in antiquity as different cereals and they were even called different names. Yet, one has to bear in mind that hulled and naked wheat forms can belong to the same biological species (Table 3). In bread wheat, T. aestivum, the difference between hulled and free-threshing varieties is governed mainly by a single mutation (the q gene). In contrast, in most free-threshing, T. turgidum wheats, the

CEREALS

25

E

A

D upper B lower

C

F

Fig. 3 Diploid einkorn wheat, Triticum monococcum. Left: A–ear (1:1), B–shattering spikelet (2:1), upper–upper disarticulation scar, lower–lower disarticulation scar, and C–grain (3:1) of wild einkorn, T. monococcum subsp. baeoticum. Right: D–ear (1:1), E–non-shattering spikelet (2:1), and F–grain (3:1) of domesticated einkorn, T. monococcum subsp. monococcum (Schiemann 1948).

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DOMESTICATION OF PLANTS IN THE OLD WORLD

B

A D

C

E

Fig. 4 Tetraploid hulled emmer wheats, Triticum turgidum. Left: A–ear (1:1), B–spikelet (2:1), and C–grain (3:1) of wild emmer wheat, T. turgidum subsp dicoccoides. Right: D–ear (1:1), and E–grain (3:1) of domesticated emmer wheat, T. turgidum subsp. dicoccum (Schiemann 1948). Note the smooth disarticulation scars in B, both below and above the internode, as in Fig. 3B.

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27

A C

B

D

Fig. 5 Tetraploid free-threshing emmer wheats, Triticum turgidum. Left: A–ear (1:1) and B–grain (3:1) of free-threshing durum wheat, T. turgidum subsp. durum. Right: C–ear (1:1), and D–grain (3:1) of free-threshing rivet wheat, T. turgidum subsp. turgidum (Schiemann 1948).

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DOMESTICATION OF PLANTS IN THE OLD WORLD

A B

C

D

Fig. 6 Hexaploid bread wheats, Triticum aestivum. A–Ear and grain of spelt wheat, T. aestivum subsp. spelta. B–Ear and grain of club wheat, T. aestivum subsp. compactum. C and D–Ears and grains of awned and awnless varieties of bread wheat, T. aestivum subsp. aestivum. Ears 1:1; grains 3:1 (Schiemann 1948).

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29

Table 3 Taxonomic classification of wheats, Triticum L. Morphological types (or species) according to traditional classification and their modern grouping on the basis of cytogenetic and molecular affinities (Van Slageren 1994). Dashed lines separate between ploidy levels. Modern grouping (biological species) (i) Diploid (2n = 14) einkorn wheat Genomic constitution: AA Both wild and domesticated forms. Collective name: T. monococcum L 1. T. monococcum L. subsp. aegilopoides (Link) Thell.

2. T. monococcum L. subsp. monococcum (ii) Diploid (2n = 14). Genomic constitution: AA 1. T. urartu Tuman ex Gand. (iii) Tetraploid (2n = 28) wild and domesticated emmer wheats, durum-type wheats, etc. Genomic constitution: BBAA Both wild and domesticated forms Collective name: T. turgidum L. 1. T. turgidum ssp. dicoccoides (Körn. ex Asch. & Graebn.) Thell. 2. T. turgidum L. ssp. dicoccum (Schrank) Thell.* 3. T. turgidum L. ssp. durum (Schrank) Thell. 4. T. turgidum L. ssp. turgidum 5. T. turgidum L. ssp. polonicum (L.) Thell. 6. T. turgidum L. ssp. carthlicum (Nevski) Löve & Löve 7. T. turgidum L. ssp. parvicoccum Kislev (iv) Tetraploid (2n = 28) Timopheev’s wheat Genomic constitution: GGAA Both wild and domesticated forms Collective name: T. timopheevii Zhuk. 1. T. timopheevii Zhuk. ssp. armeniacum (Jakubz.) van Slageren 2. T. timopheevii Zhuk. ssp. timopheevii (v) Hexaploid (2n = 42) bread wheat Genomic constitution: BBAADD (the D genome contributor is Aegilops tauschii) Only domesticated forms Collective name: T. aestivum L. 1. T. aestivum L. ssp. spelta (L.) Thell. 2. T. aestivum L. ssp. macha (Dek. & Men.) MK 3. 4. T. aestivum L. ssp. aestivum 5. T. aestivum L. ssp. compactum (Host) MK 6. T. aestivum L. ssp. sphaerococcum (Percival) Mk (vi) Hexaploid (2n = 42) Zhukovsky’s wheat Genomic constitution: GGAAAA Only domesticated forms 1. Triticum zhukovskyi Men. & Er. (non-brittle, hulled)

Traditional classification

1. Wild einkorn T. baeoticum Boiss. emend. Schiem. (brittle, hulled). Including single-grain forms (subsp. aegilopoides) and two-grain forms (subsp. thaoudar) 2. Domesticated einkorn T. monococcum L. (non-brittle, hulled) Only wild forms 1. Wild T. urartu Tuman. (brittle, hulled)

1. Wild emmer T. dicoccoides (Körn. ex. Aschers. & Graebner) Schweinf. (brittle, hulled) 2. Domesticated emmer T. dicoccum Schübl. (non-brittle, hulled) 3. Macaroni or hard wheat T. durum Desf. (domesticated, free-threshing) 4. Rivet wheat T. turgidum L. (domesticated, free-threshing) 5. Polish wheat T. polonicum L. (domesticated, free-threshing) 6. T. carthlicum Nevski [= T. persicum Vav.] (domesticated, free-threshing) 7. T. parvicoccum Kislev small grained archaeobotanical forms (domesticated, free-threshing)

1. Wild Timopheev’s wheat T. araraticum Jakubz. (brittle, hulled) 2. Domesticated Timopheev’s wheat T. timopheevii Zhuk. (non-brittle, hulled) (the uncertain ‘new’ glume wheat might belong to this group)

1. Spelt T. spelta L. (non-brittle, hulled) 2. T. macha Dekr. & Men. (non-brittle, hulled) 3. T. vavilovii Tuman. (non-brittle, hulled) 4. Bread wheat T. aestivum L. [= T. vulgare Host; T. sativum Lam.] (free-threshing) 5. Club wheat T. compactum Host. [= T. aestivo-compactum Schiem.] (free-threshing) 6. Indian dwarf wheat T. sphaerococcum Perc. (free-threshing)

Note: hulled wheat = glume wheat. * In this book, we use ‘dicoccum’ rather than Van Slageren’s ‘dicoccon’, as many taxonomists believe this is the correct spelling. (https://www.ksu.edu/wgrc/Taxonomy/taxintro.html is a good source for wheat taxonomy)

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DOMESTICATION OF PLANTS IN THE OLD WORLD

shift from ‘hulledness’ to ‘nakedness’ was brought about by a polygenic system. 2. Another important morphological trait in wheats is the manner in which the ear shatters in wild forms, or stays intact in the domesticated forms. Wild wheats are adapted to disseminate their grains by having brittle ears that disarticulate at maturity into individual spikelets (dispersal units). In wild einkorn (Fig. 3), wild emmer (Fig. 4), and wild Timopheevi’s wheat, the point of disarticulation is the upper and lower abscission scars (Fig. 3, 4 and 7). Each spikelet with a wedge-shaped rachis internode at its base constitutes an arrow-like device that inserts the grain into the ground (Zohary and Brick 1961; Zohary 1969; Harlan et al. 1973). In Aegilops tauschii, and consequently in spelt wheat, the disarticulation point is frequently below the upper abscission scar. The dispersal unit is a ‘barrelshaped’ spikelet with a characteristic rachis internode joined to its upper side (Figs 7C and 14, below and p. 49). In contrast to wild types, all domesticated wheats have non-brittle ears that stay intact after maturation. Thus, they depend on the farmer for their reaping, threshing, and sowing. Yet, threshing fractures the ears of the various domestic, nonbrittle wheats in different ways, and significantly these morphological differences are also discernible in archaeobotanical remains.

The following are indications for discerning between domesticated and wild material wheats in archaeobotanical remains (see Table 3): (i) Usually, the ears of hulled diploid and tetraploid domesticated wheats (einkorn, emmer, and Timopheevi’s wheats) break during threshing at the same place at which their wild counterparts disarticulate spontaneously. In other words, threshing in these still primitive wheats ‘mimics’ the shattering pattern of their wild progenitors. The individual spikelet with the internode at its base is the product of threshing (Fig. 7, A and B, Plate 3). However, instead of the smooth abscission scars, which characterize the wild forms, the surface of the breakage scars in the domestic hulled wheats is rough. Therefore, prevalence of spikelet forks showing rough scars (Plate 3, Fig. 8) in wheat archaeobotanical assemblages, serves as a critical indication of domestication. This criterion fits most other grain crops as well. (ii) The breakage point at each spikelet of hulled hexaploid spelt-type wheats is endowed by the wild DD contributor. In spelt wheat, this diagnostic pressure-fracture breaks the internode below the disarticulation scar. Therefore, the rachis internode remains attached to the upper side of the spikelet (Fig. 7C). In archaeobotanical assemblages, spike-

Fig. 7 The threshing products of the three main types of cultivated hulled wheats: A–Einkorn, Triticum monococcum subsp. monococcum. B–Emmer, T. turgidum subsp. dicoccum. C–Spelt, T. aestivum subsp. spelta (fresh material). Note that in A and B the rachis internode is attached below the dispersal unit, while in C it is attached above it.

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lets which follow the disarticulation pattern of Ae. tauschii indicate the presence of hexaploid spelttype wheats, though this identification must be confirmed using glume shape and texture. (iii) In free-threshing domesticated wheats (tetraploid durum-type and hexaploid bread wheats) the rachis of the ear is thickened throughout its length. Threshing breaks the glumes, lemmas, and paleas at or near their base, and releases the naked grains. The thickened rachis breaks irregularly, frequently into segments of two to five internodes (Fig. 9). In the hulled wheats, burning frequently destroys the upper parts of the investing glumes and lemma/

31

palea, exposing and freeing the grains. In contrast, the ‘spikelet forks’ (rachis internode and glume bases attached above it, Fig. 8), or its separated glume bases, often survive charring as discrete units. When found in archaeobotanical assemblages, such charred units are telltale indicators for the recognition of hulled wheats. In free-threshing wheats, the glumes and pales are thin and papery, and frequently do not survive burning. Also, the glumes, pales and most of the rachis fragments are winnowed out as chaff. As a result, they are rare in archaeobotanical assemblages. In contrast, some of the fragments of the tough rachis (particularly the more basal parts) are

Fig. 8 Charred remains of domesticated einkorn wheat retrieved from a Michelsberger culture grave at Regensburg, Germany. (A): spikelet forks, showing the rough scars typical of domesticated forms. (B): grains from the same hoard (Photographs kindly provided by D. Kučan).

0

5 mm

Fig. 9A Rachis segments, the diagnostic traits for the recognition of free-threshing wheats in archaeological remains. Carbonized rachis fragments of 4X free-threshing wheats, Neolithic Tell Ramad, Syria (van Zeist 1976).

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DOMESTICATION OF PLANTS IN THE OLD WORLD

Free-threshing forms of 4× Triticum turgidum

Free-threshing forms of 6× Triticum aestivum

1 1 2

2 3 3 4

4

5 5

5

5

Fig. 9B The structure of the rachis fragment in tetraploid and in hexaploid free-threshing wheats, pointing out five diagnostic traits for distinguishing between 4X Triticum turgidum cultivars (excluding carthlicum forms) and 6X T. aestivum cultivars (Hillman 2001, and pers. comm.).

Diagnostic traits

Tetraploid free-threshing wheats

Hexaploid free-threshing wheats

1. Lower part of the glumes

Usually retained after threshing, including the lower part of the keel A prominent lump is present beneath each glume base The sides of the internodes are more or less straight, and converge gradually downwards The internode is relatively thick, smooth, with rounded edge

Deciduous and fully removed by threshing

2. Shape of the glume base 3. Shape of the rachis internode 4+5. Structure of the edge of the internode

similar to the kernels in their size and weight, and upon winnowing they occasionally stay with the grains. Such fragments usually char well and become critical markers (Fig. 9A) for the presence of free-threshing wheats in archaeological excavations (van Zeist 1976; Hillman 1984). Such rachis segments can also be used for distinguishing between tetraploid and hexaploid freethreshing wheats (Maier 1996; Hillman et al. 1996; Jacomet 2006). The diagnostic traits separating the two chromosomal levels are given in some detail in Fig. 9B

The lump is missing (or is only very rudimentarily developed) The sides of the internodes are curved The edge is thinner, grooved by a fine longitudinal furrow, with ridges frequently supporting short hairs

(above). Finds of naked forms, belonging to both T. turgidum and T. aestivum, have been lumped together in archaeobotanical reports prior to the 1990s either as T. turgidum-T. aestivum or as ‘aestivo-compactum’. However, several reports have appeared, in which the 4x turgidum and the 6x aestivum free-threshing remains have been soundly identified (for review see Maier 1996). In some naked wheat samples obtained from Neolithic lake dwellings in central Europe, such morphological identification verified the results of ancient DNA analysis (e.g. Schlumbaum et al. 1998).

Table 4 The wild progenitors of the main domesticated wheats and their principal derivatives under domestication Wild einkorn wheat (2x) T. monococcum subsp. aegilopoides, diploid (AA), brittle ears, hulled grain.

domestication

Domesticated einkorn wheat T. monococcum subsp. monococcum, diploid (AA), non-brittle ears, hulled. Widely cultivated in the past; relic today.

Wild emmer wheat (4x) T. turgidum subsp. dicoccoides, tetraploid (BBAA), brittle ears, hulled grain.

domestication

Wild Aegilops tauschii, diploid (DD), brittle ears, hulled grains. Not domesticated, but contributed its genome to hexaploid wheats. addition of the D genome through spontaneous hybridization and chromosome doubling

Hard wheats Domesticated T. turgidum, tetraploid (BBAA), nonbrittle ears. 1st stage: hulled emmer wheat T. turgidum subsp. dicoccum (widely distributed in the past; relic today). 2nd stage: free-threshing forms, mostly durum wheat. Derived spontaneously from T. turgidum subsp. dicoccum (common in Mediterranean-type climates). Spelt and bread wheats (6x) Domestiated T. aestivum, hexaploid (BBAADD), non-brittle. 1st stage: hulled spelt-type forms (relic today). 2nd stage: free-threshing bread wheat. Derived spontaneously from spelt-type (the most common and most variable wheat crop today).

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DOMESTICATION OF PLANTS IN THE OLD WORLD

Einkorn wheat: Triticum monococcum Einkorn wheat is a diploid (2n = 2x = 14 chromosomes), a relatively uniform annual grain crop, with characteristic hulled grains, and delicate ears and spikelets (Fig. 3). A single non-shattering domesticated form of this wheat (usually referred to as T. sinskajae) is apparently a free-threshing mutant. Most domesticated einkorn varieties produce one grain per spikelet, hence its name, but cultivars with two grains exist as well (Fig. 8B). Today einkorn is a relic crop, although it is still sporadically grown in western Turkey, the Balkan countries, Germany, Switzerland, Spain, as well as Caucasia (Nesbitt and Samuel 1996a). In the past, einkorn cultivation was much more extensive. This wheat was one of the founder grain crops of Neolithic agriculture in the Fertile Crescent and a principal component of the early crop assortment in Europe, especially in the Aegean region. Since the Bronze Age, its importance seems to have declined gradually, very likely due to the competition from free-threshing wheats. Einkorn is a small plant, rarely more than 70 cm high, with a relatively low yield, but it can survive on poor soils where other wheat types fail. The fine yellow flour is nutritious, but gives bread of poor rising qualities. Thus einkorn has been consumed primarily as porridge or as cooked whole grains. Since Roman times, a considerable part of the yield has been fed to animals (Schiemann 1948; Harlan 1981).

Wild ancestry The wild ancestry of domesticated einkorn wheat is well established. Domesticated Triticum monococcum subsp. monococcum is closely related to a group of wild and weedy wheat forms spread over southwest Asia and adjacent territories, and traditionally referred to as wild einkorn or T. baeoticum Boiss. emend. Schiem. (Table 3). Both wild and domesticated einkorns are morphologically similar (Fig. 3). Both are diploid and contain identical chromosomes (genomic designation AmAm). Hybrids between wild aegilopoides and domesticated monococcum are fully fertile and chromosome pairing in their meiosis is normal. The principle distinguishing trait between wild and domesticated einkorn is the mode of seed dispersal. Wild forms have brittle ears and

the individual spikelets disarticulate at maturity to disperse the seed. In domesticated einkorn this essential adaptation to wild conditions no longer exists. The mature ear remains intact and breaks into individual spikelets only upon pressure (during threshing and/or flailing). Seed dispersal depends on reaping and sowing by humans. Another diagnostic character indicating domestication is the shape of the grain (van Zeist 1976). In domesticated forms, the kernels tend to be wider compared to the wild forms (Fig. 3, p. 25), but this sign is less reliable (Gegas et al. 2010) (see p. 22). Most specialists today regard the aegilopoides wheat not as an independent species but as the wild progenitor of the domesticated crop. It is taxonomically placed as a subspecies within the crop complex as T. monococcum L. subsp. aegilopoides (Link) Thell. [= subsp. baeoticum (Boiss.) A. et D. Löve]. Wild einkorn is widely distributed over western Asia and can be found also in the southern Balkan countries (Harlan and Zohary 1966; Zohary 1969). Its distribution centre lies in the central part of the Fertile Crescent (i.e. northern Syria, southern Turkey, northern Iraq and adjacent Iran, as well as some parts of western Anatolia) (Map 3). In these areas, wild einkorn is widely distributed as a component of oak park-forests and steppe-like formations. In addition to occupying such primary habitats, wild einkorn also grows as a weed and colonizer of secondary habitats, such as edges of cultivation and roadsides. Sometimes it also invades fields of domesticated cereals. Aegilopoides wheats are distributed over a wide ecological range with respect to both soils and climates. Edaphically, wild einkorn shows a definite affinity to basaltic soils, marls, clays, and limestones. Climatically, it thrives in the summer-dry foothills of the northern Euphrates basin, as well as on the bitterly cold, elevated plateau of central and eastern Anatolia (1400–2000 m altitude), with its summer rains. However, it does not succeed in hot and very arid climates. Further away from its distribution centre, wild einkorn is restricted mainly to segetal or secondary habitats, i.e. sites that were probably not available before the opening up of these areas to agricultural activity. Several major eco-geographical and morphological forms can be recognized in wild einkorn. In the

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0 0

100 200

35

200 miles 400 km

Map 3 Geographical distribution of wild einkorn wheat, Triticum monococcum subsp. baeoticum (= T. baeoticum). The area in which wild einkorn is widely distributed is shaded. Dots represent additional sites outside the main distribution area, harboring mainly weedy forms (based on Zohary 1989a).

north and north-west parts of its range, plants with small, single-awned, one-seeded spikelets prevail. These are sometimes named T. aegilopoides (Link) Bal. In the summer-dry southern areas, more robust plants with two-grained, two-awned spikelets, sometimes referred to as T. thaoudar Reuter, are common. But in central Anatolia, Transcaucasia, and adjacent territories of Iran, a series of intermediate forms, bridging aegilopoides and thaoudar forms abound. In fact, many Anatolian populations of wild einkorn show a wide range of variation in spikelet morphology and include single-grained individuals (with a single awn), two-grained plants (with two well-developed awns) and various intergradations between the two extreme types. Heun et al. (1997) examined sixty-eight representative lines of domesticated einkorn Triticum monococcum subsp. monococcum and 268 lines of wild einkorn T. monococcum subsp. aegilopoides for ampli-

fied fragment length polymorphism (AFLP) DNA analysis. The latter originated from the ‘Fertile Crescent’ (194 samples) and from several other parts of the distribution area of this wild wheat (74 samples). Among the wild einkorn lines tested, a group of eleven lines (which came from Karacadağ range in south-east Turkey) turned out to be distinctively separated from all other wild einkorn lines. These were genetically most similar to the tested domesticated einkorn lines, indicating that they belong to the wild source from which the crop could have evolved. The data obtained support the assumption of a monophyletic origin of domesticated einkorn wheat. Recently, Kilian et al. (2007) conducted a much larger study of genetic markers of 321 wild lines and 92 domesticated lines of einkorn. This study has found that three wild einkorn races arose prior to domestication without human intervention, and only one of them became domesticated.

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DOMESTICATION OF PLANTS IN THE OLD WORLD

According to this study, this race (which was collected from two regions in Turkey, Karacadağ and the Kartal-Karadağ range) was subjected to multiple independent domestication events, possibly after the wild grains from these two areas in south-east Turkey were dispersed throughout a wider region by the earliest cultivators. The Fertile Crescent belt also harbours T. urartu Tuman. (see above), a diploid wild wheat which resembles closely, and is easily confused with, the two-grained forms of wild einkorn (Valkoun et al. 1998). However, T. urartu can be recognized by its spreading awns, the presence of a small third awn at the tip of the spikelet, the lack of hairs on the basal leaves, and by the colour of its grains (Waines and Barnhart 1992; Waines 1996). It shows a clear preference for basaltic soils, and frequently forms mixed stands with wild einkorn wheat.

Archaeological evidence Einkorn wheat was probably collected extensively from the wild before its introduction to cultivation. The earliest reported einkorn finds are the

charred, narrow, wild-type kernels (Fig. 10), from Epi-Palaeolithic pre-agriculture layers, ca. 12,700– 11,100 cal BP, Tell Abu Hureyra (Hillman 1975, 2000a; Hillman et al. 1989) and ca. 11,800–11,300 cal BP in Mureybit (van Zeist and Bakker-Heeres 1986; van Zeist and Casparie 1968), northern Syria. Aegilopoides-type einkorn remains continue to appear in some Early PPNB (Pre-Pottery Neolithic B), 10,500–10,100 cal BP, settlements in south-west Asia where there are already definite signs of wheat and barley domestication; i.e. in Tell Abu Hureyra, Syria (Hillman 1975, 2000b; de Moulins 1997, 2000; Hillman et al. 1989), ca. 10,250–9,550 cal BP Çayönü (van Zeist and de Roller 1991–2, 2003; van Zeist 1972) and 9,450–8,450 cal BP Can Hasan III (Hillman 1972), Turkey, and (ca. 10,200–10,150 cal BP) Ali Kosh, and also in later, ca. 8,350–7,750 cal BP Tepe Sabz, Iran (Helbaek 1969). In some of these sites (as well as other contemporary southwest Asian settlements) one is faced also with plumper kernels characteristic of domesticated einkorn (Fig. 11). The first definite domesticated einkorn wheat appears in two Early PPNB sites, namely Çayönü

Fig. 10 Carbonized grains of wild einkorn wheat, Triticum monococcum subsp. baeoticum, and wild rye, Secale sp. Epi-Palaeolithic Mureybit, Syria (van Zeist and Casparie 1968). Note the elongated shape of the kernels, which characterizes wild forms.

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A

0

5 mm

37

B

Fig. 11 Comparison between charred grains of wild and of domesticated einkorn wheats, Triticum monococcum. A–Wild einkorn, T. monococcum subsp. baeoticum, from pre-agriculture Mureybit, Syria (van Zeist and Casparie 1968). B–Domesticated einkorn, T. monococcum subsp. monococcum, from early Neolithic Nea Nikomedeia, Greece (van Zeist and Bottema 1971). The upper and middle domesticated grains are from one-seeded spikelets (‘spindle shaped’), while the lower grain is from two-seeded spikelet. The grains of two-seeded einkorn spikelets are not curved in ventral side, but straight and flat, and therefore look similar to those of domesticated emmer (Fig. 12). They can be distinguished by the fact that they are slenderer, still somewhat spindle shaped in dorsal view, and have a pointed apex.

(van Zeist 1972; van Zeist and de Roller 1991–2, 2003) and Cafer Höyük (de Moulins 1997), southern Turkey (Map 1). (A third possible site, Nevali Çori (Pasternak 1998), has been mooted. We await final confirmation. Tanno and Willcox (in press) re-examined its domestication status and concluded that there is no evidence of einkorn domestication.) In these sites, numerous spikelet forks were found, showing rough breakage scars—that is, the most reliable sign of domestication. These finds indicate that einkorn belongs to the small group of annual grain plants that founded agriculture in the Fertile Crescent. It seems, then, that around 10,500–10,100 cal BP domesticated einkorn appeared first in the Euphrates Valley.

From these localions, which are situated more or less within the present range of distribution of wild einkorn (Map 3), this domesticated cereal spreads further south to Middle PPNB, ca. 10,200–9,550 cal BP, Tell el Aswad (van Zeist and Bakker-Heeres 1985) near Damascus, and to ca. 9,900–9,550 cal BP Jericho (Hopf 1983). In both these Early Neolithic sites, remains of einkorn are, however, relatively few in number. This wheat appears to have been less frequently cultivated than the two other Neolithic founder cereals, namely emmer wheat and barley (Map 1). The domesticated status of einkorn wheat from ca. 10,650–9,550 cal BP Early PPNB KissonergaMylouthkia and Shillourokambos, Cyprus (Willcox

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DOMESTICATION OF PLANTS IN THE OLD WORLD

2000; Murray 2003) relies currently on grain shape, and therefore needs further confirmation by chaff remain. However, as no wild wheats grow in the island, it seems that this find represents early introduction of agriculture from elsewhere. Somewhat later, einkorn emerges as one of the principal crops in established Neolithic food production in south-west Asia. It re-appears together with emmer wheat and barley but shows a definite preference for areas with relatively cool climates. It is rare in warmer places and totally absent from hot regions such as Egypt and Lower Mesopotamia; in the IsraelJordan region, it has been found only once—in Middle PPNB Jericho (Hopf 1983). Einkorn remains continue to be found frequently in Chalcolithic, Bronze Age, and Iron Age sites in south-west Asia. Along with emmer (pp. 44–48), it is gradually replaced by the free-threshing wheats. Einkorn wheat played an important role in the early spread of Neolithic agriculture beyond its Fertile Crescent ‘core area’. Charred remains of domesticated einkorn are numerous in the first half of the ninth millennium BP contexts of the Aegean Belt. Such sites are ca. 8,700–6,800 cal BP Cape Andreas-Kastros, Cyprus (van Zeist 1981), ca. 8,650–8,400 cal BP, Knossos, Crete (Sarpaki 2009), and ca. 8,650–8,200 cal BP, Sesklo, Greece (Hopf, 1962; Kroll, 1981a). It was also found in several early Greek and Macedonian agricultural settlements which developed during the second half of the ninth millennium BP (Renfrew 1979; Kroll 1981a), such as ca. 8,200–6,650 cal BP, Anza, Macedonia (Renfrew 1979). Recently, Valamoti (2011) concluded that einkorn dominates many Neolithic assemblages in Greece. These settlements extended as far as ca. 8,150–7,000 cal BP Obre, Bosnia-Hercegovina, former Yugoslavia (Renfrew 1974). At the same time, its reach was as far east as ca. 8,200–7,850 cal BP Jeitun, south Turkmenistan, and served there as the principal staple (Charles and Hillman 1992; Harris and Gosden 1996). Much richer remains of einkorn wheat are available from sites representing earliest agriculture in the first half of the eighth millennium BP in Bulgaria, like ca. 7,950–7,650 cal BP Early Neolithic I Kapitan Dimitrievo (Marinova 2006, forthcoming), and contemporaneous Karanovo (Thanheiser 1997; Marinova 2004, 2006) and Kovacevo (Popova 1992;

Marinova 2006). In these sites, einkorn wheat continue to be represented in large quantities up until the Bronze Age. Here (and see details in Chapter 10 and Map 1, p. 2), einkorn often prevails over emmer in frequency of pure hoards (see Thanheiser 1997; Marinova 2004, 2006, forthcoming); and retains its principal role throughout the Neolithic and Bronze Age (Hopf 1973a; Renfrew 1979). At the same time einkorn reaches ca. 7,950–7,250 cal BP StarčevoKörös culture sites in southern Hungary (Hartyányi and Nováki 1971; Hartyányi et al. 1968; Füzes 1990), as well as ca. 7,600–7,500 cal BP Sacarovca, Moldavia (Januševič 1984; Kuzminova et al. 1998; Monah 2007b). Einkorn is also one of the main cereal crops of second half of the eighth millennium Linearbandkeramik culture; i.e. the first farming settlements in central Europe (Willerding 1980; Nesbitt and Samuel 1996a). During this short period, farming spread across central and northern Europe to Spain, as well as to Egypt (where no einkorn was found). Einkorn usually occurs in a mixture with emmer wheat or sometimes as a pure crop. In some locations both cereals are equally common, but occasionally einkorn prevails. This situation persists all over Neolithic central and northern Europe, although barley becomes more frequent, and partly replaces the wheats in the later sites. Similar dominance of einkorn and emmer is found in eastern Europe, but the relative proportions of the two wheats vary considerably according to local conditions and cultures. The numerous data already available indicate that wheat cultivation in the Neolithic and Bronze Age of central Europe was mainly dependent on hulled cultivars, which do not seem to be replaced here in most cases by free-threshing wheats, as was the case in the Mediterranean basin and south-west Asia. Representative sites for this period are the Brześć group (Bieniek 2007) and contemporaneous Gwoździec, Poland (Bieniek and Lityńska-Zając 2001; Lityńska-Zając 2007); Neolithic Vinča culture, Liubcova, Rumania (Cârciumaru 1996); ZánkaVasútállomás, Hungary (Füzes 1990, 1991); Blatné, Slovakia (Hajnalová 1989); Bietigheim-Bissingen (Piening 1989) and Hienheim (Bakels 1978), Germany; Schletz, Austria (Schneider 1994; Kohler-Schneider 2007); Cova de Can Sadurní, Spain (Blasco et al. 1999); and Aubechies, Belgium (Bakels and Rousselle 1985).

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Compared to its principal role in the Balkans and in the central European Linearbandkeramik culture, einkorn is much scarcer in plant remains obtained from the early Impressed Ware culture in the west Mediterranean basin. Einkorn appears together with emmer and free-threshing wheat in the Impressed Ware (Cardial) settlements in Italy, like ca. 8,000–7,600 cal BP Scamuso (Costantini et al. 1997), ca. 7,750–7,250 cal BP Pienza (Castelletti 1976), and ca. 7,750–7,150 cal BP La Marmotta (Rottoli 1993, 2002), but rarely as a common crop. In Early Neolithic Sammardenchia and other adjacent northern Italian sites (Rottoli 1993, 2002, 2005; Rottoli and Pessina 2007), einkorn takes a more important role. It appears in Impressed Ware Early Neolithic of southern coastal France, like ca. 7,900– 7,500 cal BP Pont de Roque Haute, (Marinval 2007), and ca. 7,650–6,700 cal BP Châteauneuf-lesMartigues (Rottoli 2005; Rottoli and Pessina 2007). In Spain, einkorn arrives only in the second half of the eighth millennium, sites like ca. 7,400–7,050 cal BP Coveta de l’Or (Hopf and Schubart 1965; Lopez,1980), and adjacent Cova de Cendres (Buxó 1997), and contemporaneous Cova de Can Sadurní (Blasco et al. 1999; López et al. 2003; Peña-Chocarro 2007) and North Meseta sites (Stika 1999). In most of these contexts it does not serve as a major cereal, and it occurs afterwards in these areas only sporadically. Some einkorn also appears in early Neolithic sites in ca. 7,950–7,150 cal BP Aratashen and Aknashen, Armenia (Hovsepyan and Willcox 2008), and contemporaneous Arukhlo 1 and 2, Georgia (Lisitsina 1984; Januševič 1984; Schultze-Motel 1988a). In Transcaucasia, Chokh, Dagestan, it appears only in the end of eighth millennium BP (Lisitsina 1984), again as a member of the south-west Asian crop ensemble. The final movements of einkorn, and other domesticated crops, further into Europe is less rapid. During the seventh millennium, it reached Switzerland in ca. 6,450–5,450 cal BP Middle Neolithic sites in Sion (Martin et al. 2008a, 2008b); (Lundström-Baudais in press) and later, ca. 6,250– 4,450 cal BP Late Neolithic Egolzwil 3 (Bollinger 1994; Jacomet 2007) and Zürich (Jacomet 1988, 2006; Jacomet et al. 1989; Brombacher 1997; Favre 2002; Brombacher et al. 2005); Early Neolithic, ca.

39

6,300–5,900 cal BP, Swifterbant S3, Netherlands (van Zeist and Palfenier-Vegter 1981); and in ca. 6,000–4,750 cal BP to Eneolithic settlements in Ukraine, like Luca Vrublevecaja (Januševič 1976), Majdaneckoe, Majaki, and Usatovo (Pashkevich 2005). In the sixth millennium it reached early Neolithic Britain, ca. 5,600–5,300 cal BP The Stumble (Murphy 1989), and later to Middle Neolithic, ca. 5,000 cal BP, Alvastra, Sweden (Göransson et al. 1995; Hjelmqvist 1955) and Funnel Beaker culture, ca. 4,950–4,850 cal BP Sarup, Denmark (Jørgensen 1981). Einkorn wheat persisted in many parts of Europe until medieval times. Its cultivation almost disappeared in the twentieth and the twenty-first centuries.

Emmer and durum-type wheats: Triticum turgidum This is a varied aggregate of domesticated tetraploid (2n = 2x = 28 chromosomes) wheats. All share BBAA chromosomes, and all are almost fully interfertile with one another. Therefore, these wheats are included in a single biological species (Table 3). According to their response to threshing, the domestic turgidum wheats fall into two groups of cultivars that are frequently recognizable also in archaeological remains: (i) The products of threshing of hulled nonshattering emmer wheat, T. turgidum L. subsp. dicoccum (Schrank) Thell. (traditionally called T. dicoccum Schübl.) are the individual spikelets (Fig. 4B, Plate 3). The grains remain invested by the pales and glumes. In domestic emmer, as in einkorn, threshing results in breaking the rachis of the ear in its weakest points below each spikelet. This parallels the disarticulation pattern found in wild emmer. Taxonomically, emmer represents the primitive situation in domesticated turgidum wheats. (ii) The more advanced free-threshing tetraploids (Fig. 5) evolved under domestication from hulled emmer. The common representative of this group today is durum wheat, T. turgidum subsp. durum (Desf.) Thell. [=T. durum Desf.]. Less common types are: rivet wheat, T. turgidum subsp. turgidum L. [=T. turgidum L.]; Polish wheat, T. turgidum subsp. polon-

40

DOMESTICATION OF PLANTS IN THE OLD WORLD

icum (L.) [=T. polonicum L.]; and T. turgidum subsp. carthlicum (Nevsky) Löve & Löve [=T. carthlicum Nevski]. The glumes in all these domesticated tetraploid forms are relatively thin. Threshing breaks the glumes at their bases and frees the naked grains. The rachis is usually uniformly tough and threshing breaks it into irregular fragments. Hulled emmer was the principal wheat of Old World agriculture in Neolithic and early Bronze Age at the Mediterranean basin, south-west Asia, and temperate Europe. It was used for food and apparently also for brewing beer. Later, more advanced, naked free-threshing tetraploid and hexaploid domestic types gradually replaced it. At present, hulled emmer is still a relic crop, sporadically grown in some parts of Europe and south-west Asia; i.e. the Balkan countries, eastern Slovakia, Hungary, Spain, Anatolia, Iran, Caucasia, and India (Nesbitt and Samuel 1996b; Perrino et al. 1996). Free-threshing durum-type hard wheats are the main contemporary representatives of the BBAA tetraploid wheats. Such naked, free-threshing tetraploids (possibly including the small grain forms described by Kislev (1980, 2009) as T. parvicoccum) probably appeared already in Neolithic times and gradually gained prominence. Since classical times, free-threshing tetraploid cultivars constituted the main wheat crop in the summer-dry, relatively warm Mediterranean basin. In more continental climates and in temperate areas with summer rains, traditional wheat production depended heavily on another free-threshing wheat, namely hexaploid T. aestivum.

Wild ancestry Genetic and morphological evidence indicates (for review see Zohary 1969; Feldman et al. 1995; Feldman and Kislev 2007) that the domesticated tetraploid turgidum wheats (both the hulled dicoccum forms and the free-threshing durum-type varieties) are closely related to a wild wheat native to south-west Asia, and traditionally called T. dicoccoides (Körn. ex Aschers. et Gräbn.) Schweinfort Thell. and currently classified as T. turgidum subsp. dicoccoides (Körn. ex Asch. & Graebn.) Thell. (wild emmer wheat, see Plate 1, Plate 2, Fig. 4, Table 3). This is an annual,

predominantly self-pollinated, tetraploid wheat, with characteristic large and brittle ears and big elongated grains that show a striking similarity to some domesticated emmer (Fig. 5) and durum (Fig. 5) cultivars. It is the only wild stock in the genus Triticum that is cross-compatible and fully interfertile with the domesticated turgidum wheats. Hybrids between wild dicoccoides forms and all domesticated members of T. turgidum aggregate show normal chromosome pairing in meiosis, indicating that all these tetraploid wheats contain homologous chromosomes (BBAA chromosome sets). The close genetic affinities between wild emmer and the domesticated members of T. turgidum are also indicated by spontaneous hybridization that occurs occasionally when these wild and domesticated wheats grow side by side. On the basis of these close genetic and morphological relationships, wheat researchers regard subsp. dicoccoides as the wild progenitor of the domesticated tetraploid emmer and durum wheats. They rank wild emmer as the wild subspecies of the T. turgidum crop complex (see Tables 3 and 4). In the tetraploid turgidum wheats, the most conspicuous diagnostic difference between wild and tame is the seed-dispersal mechanism (Zohary and Brick 1961; Zohary 1969; Elbaum et al. 2007). Wild dicoccoides wheats have brittle ears that shatter upon maturity into individual spikelets. Each spikelet operates as an arrow-like device disseminating the seed by inserting them into the ground. The ‘wildtype’ rachis disarticulation and the spikelet morphology reflect specialization in seed dissemination. This ensures the survival of the wild forms under wild conditions. Under the manmade system of tillage, reaping, threshing, and sowing, this adaptation broke down and non-brittle types were automatically selected. Significantly (Table 7, p. 61), the shift from a brittle spike (in wild dicoccoides wheat) to a non-brittle spike (in domesticated dicoccum wheats) is governed by recessive alleles at two major loci that are located respectively on the short arm of chromosomes 3A and 3B (Watanabe et al. 2002, 2006; Nalam et al. 2006). In addition, wild and domesticated forms differ from one another in kernel morphology (van Zeist 1976). In domesticated dicoccum and durum forms, the grain tends to be wider and thicker as well as rounder in cross-section com-

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pared to the wild dicoccoides counterpart (compare grains in Figs 4 and 5). These domestication traits are quite helpful in analyzing archaeological remains. Wild emmer, T. turgidum spp. dicoccoides (Körn. ex Asch. & Graebn.) Thell. [=T. dicoccoides (Körn.) Schweinf.] is more restricted in its distribution and more confined in its ecology than wild einkorn. Its range covers the Fertile Crescent: Israel, Jordan, south-western Syria, Lebanon, south-eastern Turkey, northern Iraq, and western Iran (Map 4). It is most widespread in the catchment area of the upper Jordan Valley. Wild emmer was found in 1906 in eastern Galilee and on the slopes of Mt Hermon in Israel (Aaronsohn 1910; Schiemann 1956). Wild dicoccoides wheats grow as common annual components in the herbaceous cover of the Tabor oak (Quercus ithaburensis) park-forest belt and related steppe-like herbaceous plant formations. They are confined to basaltic and hard limestone bedrocks and are completely absent on marls and chalks. In rocky places that have not been severely overgrazed, dicoccoides wheat often grows in large stands. With wild barley Hordeum spontaneum and wild oat Avena sterilis, they form ‘fields of wild cereals’. Further north, wild emmer occurs in the AntiLebanon range and again in the oak park-forest belt of Quercus brantii in south-eastern Turkey, north Iraq, and west Iran (Map 4). Whereas dicoccoides wheat occurs alone in the Levant, it grows sympatrically with a second wild tetraploid wheat, T. timopheevi subsp. armeniacum (Jakubz.) van Slageren (see p. 51), in the north-eastern part of its distribution area. Cytogenetically, these two wild wheats have distinct genomic and cytoplasmic constitutions and are separated from one another by strong sterility reproductive barriers (Sachs 1953; Wagenaar 1966; Maan 1973). However, they are so similar morphologically that it is very hard or even impossible to separate between ears of the two species without cytogenetic confirmation or the use of molecular genetic markers. In the north and the north-east, dicoccoides wheat is closely associated not only with Hordeum spontaneum but also with wild einkorn, T. monococcum subsp. baeoticum and T. urartu. In north Israel and in south Syria, dicoccoides wheat shows a multitude of forms and often builds con-

41

spicuously polymorphic populations that are easily noticed by their variation in glumes’ hairiness, colour of the spike, the size of the spikelets, and the shape of the leaves (Poyarkova 1988; Poyarkova and Gerechter-Amitai 1991). Climatically too, wild emmer shows a considerable range of adaptation. It is distributed over a wide altitudinal range. Robust, early maturing, types occupy the winter-warm basin around the Sea of Galilee to altitudes as low as 100 m below sea level. More slender, late-blooming forms occur higher up in the Galilee mountains and climb to elevations of 1600 m on the east- and south-facing slopes of Mt Hermon. In the Zagros Mountains of north Iraq and west Iran, dicoccoides wheats occur at altitudes ranging from 700 to 1600 m. Molecular studies have placed a probable single origin of emmer wheat in Turkey. Özkan et al. (2002) examined twenty-four AFLP loci of ninety-noine wild emmer lines from all over the Fertile Crescent belt and several dozens of different cultivars. They showed that the most likely place of origin for emmer wheat is in south-eastern Turkey. Recently, a study based on 131 RFLP loci, Luo et al. (2007) reached similar results.

Archaeological evidence (a) Hulled emmer wheats: Similar to einkorn wheat (p. 34) and barley (p. 51), emmer wheat was collected from the wild long before its domestication. Brittle, dicoccoides-like remains, with relatively narrow grains (Fig. 12) appear in several early Fertile Crescent sites. A clear indication of preagriculture gathering of wild emmer wheat is available from the south-western shore of the Sea of Galilee in Israel. Here, remains of subsp. dicoccoides (as well as wild barley) were recovered from Ohalo II, a submerged ca. 23,000 cal BP early EpiPaleolithic site (Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004, 2008). Dicoccoidestype grains were discovered in ca. 11,700–10,550 cal BP, PPNA Netiv Hagdud and Gilgal, Israel (Kislev 1997; Hartmann 2006; Weiss et al. 2006). The earliest fully convincing sign to-date of domesticated emmer come from the numerous spikelet forks retrieved from ca. 10,250–9,550 cal BP Early PPNB Çayönü (van Zeist 1972; van Zeist and de

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DOMESTICATION OF PLANTS IN THE OLD WORLD

0 0

100 200

200 miles 400 km

Triticum dicoccoides Triticum ararticum

Map 4 Geographical distribution of wild emmer wheat, Triticum turgidum subsp. dicoccoides [= T. dicoccoides] and wild Timopheev’s wheat, T. timopheevii subsp. araraticum [= T. araraticum], that grow side by side in the same habitats (compiled from Harlan and Zohary 1966; Rao and Smith 1968; Dagan and Zohary 1970; Maan 1973; Tanaka and Ishii 1973; Tanaka et al. 1979; and unpublished data of D. Zohary).

Roller 1991–2, 2003). Here, hundreds of spikelet forks were discovered, all showing rough breakage scars characteristic of domestic emmer. A rough breakage scar is by far the best indication that we are dealing with domesticated forms of wheat in archaeological deposits. These spikelet forks are accompanied by wild-type narrow grains, which give some indication that mutant rachises predated the increase in seed-size in the domestication process. Numerous spikelet forks, with similar telltale rough scars, are available also from contemporary contexts at Cafer Höyük (de Moulins 1997). (A third possible site, Nevali Çori (Pasternak 1998), has been announced only preliminarily; we wait for its final publication.) All these finds indicate that at Early PPNB, emmer domestication must have been well underway in the Fertile Crescent. The domesticated status of emmer wheat from ca. 10,650–9,550 cal BP Early PPNB Kissonerga-

Mylouthkia and Shillourokambos, Cyprus (Willcox 2000; Murray 2003) was not confirmed yet by chaff remains. Even so, as no wild wheats grow in the island, it seems that this find represents early introduction of agriculture from outside. A less reliable diagnostic trait for early domestication of wheat is grain size. In cultivars, seeds are usually plumper and larger than those in their wild relatives (see grain sizes in Figs 3–4, 11–12). However, this process takes longer, and therefore size increase cannot indicate reliably the start of domestication (see p. 22). To-date, the earliest conspicuous grain-size increase is reported from Early PPNB Tell Aswad, 25 km south-east of Damascus (van Zeist and Bakker-Heeres 1985). Here, dicoccum-like plump kernels start to appear in the lowest habitation level Ia, 10,500–10,200 cal BP (van Zeist and BakkerHeeres 1985; Willcox 2005), and also in Middle

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A

0

5 mm

43

B

Fig. 12 Comparison between carbonized grains of wild and of domesticated emmer wheats, Triticum turgidum, Neolithic Çayönü, Turkey (van Zeist 1972). A–Wild emmer wheat, Triticum turgidum subsp. dicoccoides [= T. dicoccoides]. B–Domesticated emmer wheat, Triticum turgidum subsp. dicoccum [= T. dicoccum]. Note that compared to the more ‘spindle-shaped’ grains of one-seeded spikelets of domesticated einkorn (Fig. 13) the grains of cultivated emmer have a wider, blunter apex and a straighter side shape.

PPNB phases II (10,200–9,550 cal BP). In these phases, however, no rachis segments have been retrieved. Significantly, no wild dicoccoides-like narrow kernels were retrieved from Tell Aswad. The present climate in the rain-shadowed Damascus basin is far too dry for wild wheat, and it was probably arid 10,000 years ago also). As van Zeist and Bakker-Heeres emphasize, the continuous presence of morphologically discernible dicoccum kernels, the total absence (from the very start) of dicoccoides-like material, and the extreme dryness (less than 200 mm annual rainfall) suggest that emmer wheat was introduced into the Damascus basin as a domesticated cereal, not later than thesecond half of the eleventh millennium BP. From ca. 9,800–8,700 cal BP onward, charred grains, that morphologically conform with dicoccum, appear also at Tell Abu Hureyra, north-east Syria, again with no rachis seg-

ments (Hillman 1975, 2000b; Hillman et al. 1989), and in contemporary Pre-Pottery Neolithic B ca. 9,450–8,450 cal BP, Can Hasan III and ca. 9,350–8,950 cal BP, Çatalhöyük East, Turkey (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007), ca. 9,600–8,750 cal BP Ali Kosh, Khuzistan (Helbaek 1969), ca. 9,900– 9,550 cal BP, Jericho, Israel (Hopf 1983) and ca. 9,450–9,300 cal BP, Jarmo, Iraq (Braidwood 1960; Helbaek 1959a, 1960). From the very beginnings of agriculture in the Fertile Crescent (some 10,500–10,100 cal BP), emmer is the principal cereal of the newly established PrePottery Neolithic B (PPNB) farming settlements (Map 1). Although remains of domesticated einkorn and domesticated barley also occur quite regularly in these contexts, emmer prevails quantitatively in most cases. At several of the examined sites the remains of emmer include not only carbonized

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DOMESTICATION OF PLANTS IN THE OLD WORLD

plump seeds, but also well-preserved spikelets or spikelet forks showing the effects of pressure fracturing at the base of the rachis internode (Figs 7 and 8). The prevalence of rough scars indicates that this wheat had non-brittle ears. This is unequivocal proof that there and then, grain cultivation was already in practice. Emmer wheat continues to be the principal grain crop during the later stages of the Neolithic in south-west Asia, and it was apparently widely grown in this area also in Chalcolithic and Bronze Age times, although in the later periods the hulled cultivars (emmer and einkorn) were replaced by free-threshing wheats. Emmer wheat was also the main crop in the spread of the Neolithic agricultural technology from the Fertile Crescent ‘core area’ towards the west, the north, the east, and the south. It is the most common constituent of the crop assemblage that started agriculture in the Aegean region in the ninth millennium BP. Such sites are ca. 9,000–7,600 cal BP, Dhali Agridhi, Cyprus (Stewart 1974), ca. 9,000– 8,050 cal BP, Franchthi Cave (Hansen 1991b, 1992), and ca. 8,650–8,200 cal BP, Sesklo, Greece (Hopf 1962; Kroll 1981a), and ca. 8,650–8,400 cal BP, Knossos, Crete (Sarpaki 2009) (for review, see Renfrew 1979; Kroll 1981a, 1981b, 1991). In the second half of the ninth millennium BP, emmer wheat is continuing to spread to the Balkan countries, such as ca. 8,150–7,000 cal BP, Obre, Macedonia (Renfrew 1974, 1979; Kroll 1991). In the early Neolithic, it also spread to Central Asia ca. 8,200–7,850 cal BP, Jeitun (Djeitun) (Charles and Hillman 1992; Harris and Gosden 1996). Emmer was the principal cereal of the Linearbandkeramik farmers that started Neolithic agriculture in central Europe in the first and second half of the eighth millennium BP. Evidence can be found at sites like ca. 7,950–5,250 cal BP, KörösStarčevo, Hungary (Hartyányi et al. 1968; Hartyáni and Nováki 1971, 1975; Füzes 1990); ca. 7,650–7,400 cal BP Mohelnice, Czech Republic (Opravil 1979, 1981; Kühn 1981); ca. 7,500–6,550 cal BP, Aldenhovener Platte, Germany (Knörzer 1973, 1974, 1997); ca. 7,450–6,950 cal BP, Brześć, Poland (Bieniek 2007); ca. 7,150–6,800 cal BP, Schletz, Austria (Schneider 1994; Kohler-Schneider 2007). At the numerous Linearbandkeramik sites exam-

ined to-date (for review, see Willerding 1980; Nesbitt and Samuel 1996b), emmer is usually found side by side or in admixture with einkorn. Yet in most places, emmer is the more common wheat. This relationship persists all over central and northern Europe during the later Neolithic and the Bronze Age. Thus in central Europe, wheat production during the Neolithic and Bronze Age depended heavily on hulled wheats. Naked wheats do not seem to have replaced them as quickly as they did in the Mediterranean basin and in south-west Asia, though they are present in central Europe since the Neolithic. Rich finds of emmer have also been discovered at places belonging to the Impressed Ware culture that introduced agriculture into the western Mediterranean basin during the eighth millennium BP (Hopf 1991). Such sites include ca. 8,000–7,600 cal BP, Scamuso, Italy, ca. 7,400–7,050 cal BP Coveta de l’Or and Cova de Cendres (Costantini et al. 1997), and ca. 7,400–6,650 cal BP Cova de Can Sadurní (Blasco et al. 1999), Spain. In Impressed Ware contexts, emmer is generally accompanied by free-threshing wheats, and in many localities the latter prevail. Emmer is also found in younger sites of this region, but again, only in minor quantities compared to the more advanced free-threshing wheats. Emmer is, however, the wheat of Neolithic and Bronze Age Egypt (Fig. 13, Plate 5), such as ca. 7,500–6,650 cal BP Fayum and 1,325 BC Tutankhamun tomb (Caton-Thompson and Gardner 1934; Täckholm 1976; Germer 1989a; Hepper 1990; Wetterstrom 1993; Wendrich and Cappers 2005; de Vartavan et al. 2010; see also Fig. 8). As to the diffusion of Neolithic agriculture from the Fertile Crescent towards the east, there is much less information at hand. Judging by the rare data available, emmer wheat was one of the main elements in the crop assemblage discovered in several Neolithic sites in Caucasia and Transcaucasia, dated into the eighth millennium BP, as attested by the finds from ca. 7,950–7,150 cal BP Aratashen and Aknashen, Armenia (Januševič 1984; Wasylikowa et al. 1991; Hovsepyan and Willcox 2008). Finally, it should be highlighted that among the three main Neolithic grain crops (einkorn, emmer, and barley), the wild progenitor of emmer has the

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0

45

1 cm

Fig. 13 Desiccated spikelets of domesticated emmer wheat, Triticum turgidum subsp. dicoccum (G. Schweinfurth collection, fifth-dynasty Abusir, Egypt).

most limited geographic distribution. It is confined to the Fertile Crescent ‘arc’—compare Map 3 with Maps 4 and 6. Therefore, the numerous finds of domesticated emmer retrieved from sites outside the Fertile Crescent serve as convincing evidence for the fertile crescent as the core area where Neolithic agriculture could have evolved, and subsequently disseminated. (b) Free-threshing tetraploid wheats: Free-threshing wheat forms, identifiable by their rachis fragments (Fig. 9A), made their appearance in south-west Asia soon after the appearance of domesticated emmer wheat (for review see Maier 1996). Some remains start to appear already among the plant remains of two Middle PPNB sites, ca. 10,200–9,550 cal BP Tell Aswad, Syria (van Zeist and Bakker-Heeres 1985), and ca. 10,100–9,450 cal BP Aşikli Höyük, Turkey (van Zeist and de Roller 1995). More were found

somewhat later, in ca. 9,450–8,450 cal BP Late/Final Neolithic Can Hasan III, Turkey, where numerous kernels and rachis fragments were found (Hillmam 1972, 1978), and Late PPNB 9,450–8,600 cal BP Tell Bouqras, Syria (van Zeist and Waterbolk-van Rooijen 1985). They also occur in ca. 9,350–8,950 cal BP Middle/Late PPNB Çatalhöyük, Turkey (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007), and Tell Ramad, Syria (van Zeist and Bakker-Heeres 1985). In the first stages of research, archaeobotanists could not distinguish between tetraploid and hexaploid free-threshing wheats. Almost always, they have referred to them as T. turgidum-T. aestivum or as ‘aestivo-compactum’ wheats. Yet most researchers, when faced with remains obtained from the PPNB farming villages in the Fertile Crescent, assumed that they represent 4x durum-like forms rather than 6x aestivum wheats. As argued by Zohary (1969b)

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DOMESTICATION OF PLANTS IN THE OLD WORLD

and by van Zeist (1976), hexaploid bread wheats must have originated outside the ‘Fertile Crescent area’; and this development occurred only after Neolithic wheat agriculture reached the primary habitats of Aegilops tauschii distribution area (Map 5). This contact very likely happened in the Caspian belt somewhere between 8,000 and 7,000 BP. Thus, the early naked wheats in the Fertile Crescent core area could not possibly have been 6x T. aestivum. They should be attributed to 4x T. turgidum. Nesbitt (2001) claim, in contrast, for a different scenario, based on the hexaploid rachis remains from ca. 9,450–8,450 cal BP PPNB Can Hasan III (Hillmam 1972, 1978) and from Cafer Höyük (de Moulins 1997). According to Nesbitt, these finds demonstrate either that agriculture reached the Caspian belt earlier, or that the prehistoric primary distribution of Ae. tauschii extended further west. The chromosomal level of the free-threshing wheats retrieved from later south-west Asian contexts, and from sites representing the early stages of the agricultural expansion to Europe and west Asia, has been even more problematic. In many parts of

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this vast area (for details see Maier 1996), naked wheats continue to appear—in Neolithic and in Bronze Age times—in progressively increasing proportions. But assigning them to either 4x T. turgidum or to 6x T. aestivum was until recently practically impossible. They were almost always referred to as ‘aestivo-compactum’ or T. turgidum-T. aestivum forms. From the beginning of the seventh millennium BP onward, free-threshing wheats appear in Greek sites. Further west, naked wheats abound in the Impressed Ware (Cardial) sites of Italy, south France, and Spain; i.e. in settlements which represent the early stages of agriculture expansion into the west Mediterranean basin (Hopf 1991; Maier 1996). Freethreshing wheats also abound in the Middle Neolithic lake-shore sites of Switzerland and its neighboring areas (Jacomet et al. 1991; Maier 1996; Jacomet 2004, 2006, 2007). Thus, in south-west Asia, the Mediterranean basin and the Alps, we are faced with a rather early, partial replacement of hulled wheats by free-threshing types. Emmer and einkorn continued to occur and they are occasionally retrieved in large quantities. Yet towards the Late

200 miles 400 km

Map 5 Geographical distribution of Aegilops tauschii [= Ae. squarrosa]. The areas in which primary habitats occur are shaded. Dots represent additional sites, mainly of weedy forms (based on Zohary et al. 1969; van Slageren 1994).

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Bronze Age, south-west Asia and the Mediterranean basin both commonly show a prevalence of naked wheats. The large quantities of naked wheat unearthed in Bronze Age towns in the Levant are also impressive. In contrast, the wheat of ancient Egypt was hulled emmer and it retained its dominance in the Nile Valley as late as Hellenistic times. It was replaced first by naked durum wheat by the Ptolemaic rulers (Murray 2000b; Wetterstrom 1993). Our ability to distinguish between the 4x and 6x free-threshing wheats on the basis of rachis remains is already established (Jacomet 1987; Hillman et al. 1996; Maier 1996). As noted in Fig. 9B (p. 32), a set of morphological traits, traceable in rachis segments, has been found to be diagnostically reliable for such identification. Several reports have already appeared in which the remains of naked forms of 4x T. turgidum and of 6x T. aestivum are soundly recognized. Thus Maier (1996) had convincingly demonstrated that the early free-threshing wheats grown at the Middle Neolithic lake-shore site at ca. 5,850–5,450 cal BP Hornstaad, Lake Constance, Germany, were tetraploid (Maier 2001). Maier also stresses the fact that some of the rachis segments, whole spikes, and parts of spikes retrieved from several other well-investigated, Middle Neolithic lake-shore sites at the northern foothills of the Alps, manifest 4x T. turgidum morphology (see also Jacomet et al. 1989, 1991; Brombacher 1997). These finds indicate that at that time, wheat production was already based, in part, on growing naked tetraploid forms. So far, the Alpine Middle Neolithic sites constitute the only area where the identification of the 4x naked T. turgidum (as well as 6x T. aestivum) has been carried out on a large scale. But one would expect that additional finds of ‘aestivo-compactum’ wheats, coming from other parts of Europe and west Asia, will be re-examined soon. Very likely the origins and spread of the free-threshing 4x and 6x wheats will be much better understood in a few years from now.

Bread wheat: Triticum aestivum Bread wheat is the most variable aggregate of domesticated wheats, and economically the most important wheat species. It accounts for about 90%

47

of the total world wheat production today and includes numerous and contrasting types. All varieties are hexaploid (2n = 6x = 42 chromosomes) and inter-fertile when crossed with one another. All share the BBAADD genomic constitution. Hexaploid T. aestivum is a new wheat species that evolved under domestication from the domesticated tetraploid T. turgidum stock. In contrast with the diploid and tetraploid wheats, it does not have a wild hexaploid counterpart. For cytogeneticists and plant evolutionists, bread wheat stands as one of the classic examples of evolution by polyploidy. Genome analysis studies have shown (Kihara 1944; McFadden and Sears 1944, 1946) that T. aestivum is a hybridization product between tetraploid turgidum wheat (genomic constitution BBAA) and a diploid wild grass Aegilops tauschii Coss. [= Ae. squarrosa L.] (genomic constitution DD). In other words, hexaploid bread wheats originated by the addition of a third genome to the two genomes already present in tetraploid T. turgidum. Bread wheat has been synthesized in the laboratory by crossing the two putative parents and doubling the chromosomes in the hybrids. Since no BBAADD hexaploid wheat occurs in the wild (except subsp. tibetanum which is a feral wheat that presumably escaped from cultivation), this development could have occurred only under cultivation; i.e. by the hybridization of the already domesticated tetraploid wheat with the wild Ae. tauschii. Hexaploid wheats fall into two groups according to their response to threshing: (i) All the cultivars in this group are hulled (glumed), and today they survive mainly as relic crops. Prominent among them is spelt wheat, T. aestivum subsp. spelta (L.) Thell. (traditionally called T. spelta L.). An additional type, the macha wheat, T. aestivum subsp. macha (Dekapr. et Menabde) MacKey [=T. macha Dekapr. et Menabde], endemic to western Georgia, also belongs to this group. In all hulled hexaploid wheats, the products of threshing are the individual spikelets. But while in einkorn and emmer the rachis segment below the spikelet remains attached to the unit (Fig. 7 A, B), in hulled hexaploid wheats, the segment attached is frequently the internode above the base of the spikelet (Fig. 7 C). In archaeological remains, the upper

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DOMESTICATION OF PLANTS IN THE OLD WORLD

rachis segments, as well as the characteristic structure of the glumes, serve as diagnostic traits for spelt-wheat identification. (ii) Free-threshing bread wheats (Fig. 6 A, B, C) are predominantly used today. Most important and almost universal is bread wheat or common wheat, T. aestivum subsp. aestivum MacKey [also referred to as T. aestivum L., T. sativum L., or T. vulgare Host]. Two other free-threshing hexaploid types are: (a) Club wheat, T. aestivum subsp. compactum (Host) MacKey [= T. compactum Host], with compact ears. It is grown today in Afghanistan, as well as in the north-western United States. It has also been found in early European cultures; (b) Indian dwarf wheat, T. aestivum subsp. sphaerococcum (Perc.) MacKey [= T. sphaerococcum Perc.], with characteristic small grains, is native to India and Pakistan. Experimental evidence indicates that the first hexaploid wheats were spelt-like. Artificial synthesis of T. aestivum (by crossing and fusing tetraploid BBAA T. turgidum with diploid DD Aegilops tauschii) always results in hulled products (Kerber and Rowland 1974), irrespective of whether the turgidum BBAA parent is hulled or naked. It also suggests that the free-threshing condition in hexaploid bread wheats was brought about by two events: the appearance of the free-threshing gene Q located on chromosome 5A, and the mutation from Tg to tg in the gene responsible for the tenacious glumes trait on chromosome 2D. All present-day 6x naked wheats examined carry the tg/tg Q/Q genotype. As already mentioned (pp. 32, 47), the history of the 6x free-threshing bread wheats could not be easily reconstructed first, since their archaeological remains were difficult to separate from those of 4x T. turgidum. In the past, the shape of the kernel was considered a valuable diagnostic trait, because hexaploid forms tend to have relatively plumper grains with blunt tips. However, grain shape proved to be problematic because tetraploid and hexaploid wheats overlap considerably in their seed shape. This overlap is more prominent in archaeological remains, because of swelling and other deformations caused by the charring (Hopf 1955; van Zeist 1976; Harlan 1981). Thus carbonized wheat remains showing obtuse, plump kernels need not necessarily be 6x T. aestivum. They may also represent 4x T. tur-

gidum. Similar difficulties have been encountered with another feature of the grain, namely the shape of the scutellum (the connecting tissue between the embryo and the endosperm). It is short, wide, and shows a straight profile in the bread wheats; it is elongated, curved, and shows a curved profile in the durum and einkorn wheats. Yet in this trait, one also encounters variation and some overlapping. For these reasons most archaeobotanists in the past did not attempt to distinguish between 4x and 6x naked wheats. Instead, they lumped them together as ‘aestivo-compactum’ or T. turgidum-T. aestivum finds. Not so long ago (Hillman 2001), it has become clear that free-threshing 4x T. turgidum forms can be separated from their 6x T. aestivum counterparts by the morphology of their rachis segments (Fig. 9B). Parallel to what is happening with the free-threshing 4x wheats (pp. 45–46), a more precise understanding of the history of the naked 6x wheats was suggested by Maier (1996), but more research is needed.

Wild ancestry As already stated (see above), hexaploid bread wheat originated by the addition of the DD chromosome complement of Aegilops tauschii [=Ae. squarrosa] to the domesticated, tetraploid BBAA turgidum wheats. Ae. tauschii (Fig. 14) was never domesticated as such. The examination of its ecology and distribution reveals that this wild grass contributed substantially to the adaptation and the worldwide success of the bread wheats. The first significant feature about Ae. tauschii is its distribution, which also reflects its climatic requirements (Zohary 1969). This is the easternmost diploid species in the wheat group (Triticum-Aegilops). Its centre of distribution does not lie in the Mediterranean south-west Asia but in continental or temperate central Asia. It is widespread and very common in northern Iran and adjacent Transcaucasia, Transcaspia, and Afghanistan (see Map 5, and Fig 73 in van Slageren 1994). From this geographic centre, Ae. tauschii spreads west to east Syria, and east to Pakistan. In central Asia it is recorded as far east as Kirghizia and adjacent parts of Kazakhstan. Ae. tauschii is a variable species. It is represented by a multitude of forms, from slender types with cylindrical spikes, to more robust plants with thick,

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beaded spikes (see Fig. 14). The latter (subsp. strangulata) are confined to the south Caspian basin. Ae. tauschii is not only widely distributed, it also occurs over a wide range of ecological conditions. Like wild einkorn and wild barley, it occupies both primary and segetal habitats. In the centre of its distribution this plant is a frequent annual component of open formations. It thrives in areas characterized by continental climatic conditions, from the dry sagebrush steppes of the elevated Iranian and Afghan plateaus to desert margins, as well as in more temperate climates such as the rain-soaked southern coastal plain of the Caspian Sea. At the same time, all over this area Ae. tauschii is a successful colonizer of secondary, manmade habitats and a common weed in cereal fields. Towards the periphery of its distribution, it is almost exclusively a weed in cultivation. Thus in the case of Ae. tauschii, we are faced with a colonizer plant which apparently expanded its distribution over secondary habitats with the opening up of the land by agriculture. These ecological and distributional facts provide clues to the plausible place of origin of T. aestivum and explain some of the bread wheat’s ecological specifications. At the start of agriculture, the two contributors that fused to form the hexaploid wheats were evidently geographically separated. Wild emmer was restricted to south-west Asia and Ae. tauschii did not spread westward from north Iran. Thus contacts between the tetraploid wheats and Ae. tauschii could have been established only after the domestication of emmer and the spread of domesticated tetraploid T. turgidum to north Iran and adjacent Transcaucasia. This expansion probably took place between 8,000 and 7,000 BP (van Zeist 1976). The most likely area of origin of the hexaploid bread wheat, therefore, is the south-western corner of the Caspian belt. Such association between domesticated T. turgidum and weedy Ae. tauschii in cultivation can still be found in this area (Matsuoka et al. 2008). This suggestion is supported by molecular studies which have revealed the range and the structuring of isozyme and DNA polymorphism in Ae. tauschii and compared it with its counterparts on the D genome of hexaploid T. aestivum (Dvořák et al. 1998). The assembled data indicate that populations of Ae. tauschii native to Armenia and the south-west part of the Caspian Sea belt (mainly forms taxonom-

0

1

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2 cm

Fig. 14 Lower parts of the spike, and individual spikelets, of diploid Aegilops tauschii [= Ae. squarrosa], the donor of the D genome to hexaploid Triticum aestivum wheats (from the reference collection of D. Fuller, Institute of Archaeology, UCL).

ically placed in subsp. strangulata) are closest to the genome D found in the hexaploid wheats. The addition of the D genome greatly extended the range of adaptation of domesticated wheats (Zohary 1969). As mentioned above, domesticated tetraploid wheats are fit to the Mediterranean-type environments (with mild winters and warm, rainless summers). The incorporation of the Ae. tauschii genome made the hexaploid plants more capable of withstanding continental winters and humid summers. This facilitated the spread of hexaploid bread wheats over the continental plateaus of Asia and the colder, temperate areas in eastern, central, and northern Europe. It also explains the distribution patterns of wheats in traditional agriculture; i.e. the centering of the 4x durum races in south-west Asia and the Mediterranean basin, and the prevalence of 6x aestivum forms in temperate Europe and continental west Asia. Additional explanation for the origin of European spelt as independent of the Asian spelt was already proposed in the first half of the twentieth century (e.g. Bertsch F. 1943; Bertsch K. 1950; Kuckuck and Schiemann 1957). The proponents of this hypothesis

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(a) Hulled (=glumed)spelt wheat: Spelt has been reported by Januševič (1984) from ca. 8,000–7,150 cal BP Neolithic contexts of Arukhlo 1, Transcaucasia. The finds are few, but they unmistakably show the telltale upper rachis segment attached to the lower spikelet. These data confirm several earlier, poorly documented reports by other Russian researchers (Lisitsina 1978) on the occurrence of spelt wheat in some sites in the Kura river plain, Transcaucasia, dating to the seventh millennium BP, and possibly even to the eighth millennium BP. It is therefore clear that in the seventh millennium BP, hexaploid hulled T. aestivum was already grown in the Caspian belt. Significantly these early finds come from the area most likely to be the area of formation of hexaploid wheats; i.e. a region in which the domesticated 4x T. turgidum could have first come in contact with (truly wild) 2x Ae. tauschii.

late Neolithic Poland, e.g. ca. 6,350–5,950 cal BP, Brześć (Bieniek 2007) and Rumania, e.g. ca. 6,950– 6,600 cal BP, Poduri (Cârciumaru and Monah 1985; Monah and Monah 2008), Cortaillod sur les Rochettes Est, Switzerland (Akeret 2005), and Germany (Blankenhorn and Hopf 1982). In some Tripolye culture settlements in Moldavia and Bessarabia, spelt wheat was apparently grown as a crop by itself (Januševič 1976, 1978). Several wellpreserved spikelets, admixed as an ‘impurity’ with einkorn and emmer remains, were retrieved from ca. 5,700 BP Gumelnitza culture Ovčarovo, north-eastern Bulgaria (Januševič 1978). The wellpreserved spikelets leave little doubt that hexaploid spelt wheat was already in use at that time. Spelt wheat remains from Bronze and Iron Age are more numerous and come from all over east, central, and north Europe, (Hajnalová 1978; Gyulai 2003) as well as from Greece (Kroll 1983). Spelt was also well known to the Romans (Jasny 1944) and in Europe it survived as a crop until the start of the twentieth century. (Very small quantities of spelt are still grown today in south Germany, north Spain, as well as in several other parts of Europe and west Asia). It is important to note that the presence of Neolithic spelt is questioned recently by Nesbitt (2001 and pers. comm.). In his view, all or most of these finds are either of Ae. cylindrica (a common weed), new-type emmer wheat (which has strongly striated glumes, superficially similar to those of spelt), or based on grain morphology and are thus invalid. Detailed examination of published spelt is under way and awaits publication. This view concurs with recent molecular evidence, which suggests spelt arose anew in Europe, rather than move from south-west Asia to Europe.

Another relatively early spelt wheat find comes from the ca. 7,600–7,400 cal BP Early Neolithic Sacarovca, Moldavia (Januševič 1984; Kuzminova et al. 1998; Monah 2007a). Here, only a few charred spikelets were detected among numerous remains of emmer wheat. But these clearly display the diagnostic arrangement of the rachis segment of spelt. Sometime later, spelt appears sporadically in east and central European sites, like Zánka-Vasútállomás, Hungary (Monah 2007a) and Blatné, Slovakia (Hajnalová 1989). Similar finds are available from

(b) Free-threshing bread wheat: Genetic analysis indicates (p. 48) that naked hexaploid wheats could have evolved rather quickly from their more primitive spelt relatives since the shift was apparently produced by only two mutations. Thus it is plausible to assume that shortly after the formation and establishment of hexaploid spelt, the naked derivatives could have appeared as well. In other words, from the seventh millennium BP onwards, the progressively more common free-threshing wheats encountered in archaeological digs could represent not

claim that European spelt was formed by secondary hybridization between T. aestivum and T. dicoccum (and see discussion in Nesbitt 2001). An important development for this hypothesis gain Recently, Blatter, Jacomet and Schlumbaum (2002) supported the European origin by analyzing of glutenin subunit genes B1-1 and A1–2 in fifty-eight accessions of hexa- and tetraploid wheat from Europe and Asia. Their results suggests a polyphyletic origin of the A- and B-genomes of hexaploid wheat, and that European spelt does not derive from the hulled progenitors of bread wheat but originated by introgression of a tetraploid wheat into free-threshing hexaploid wheat as a secondary evolution after the development of bread wheat.

Archaeological evidence

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only tetraploid forms but hexaploid cultivars as well. A large scale re-examination (by the discriminating rachis morphology) of early remains of ‘aestivo-compactum’ naked wheats in west Asia and Europe has not yet been attempted. The continental and temperate parts of west Asia and Europe are, ecologically, the territories best suited for the early establishment of the 6x free-threshing bread wheats. It is therefore likely that from the late Neolithic and the Bronze Age onward, remains of naked wheats from places like Caucasia, central Asia, the central Anatolian plateau, India, and east and central Europe could represent largely 6x aestivum material rather than 4x turgidum forms.

Timopheev’s wheat: Triticum timopheevii This tetraploid species comprises both domesticated and wild forms that are genomically different and reproductively isolated from the more common tetraploid turgidum stock. F1 Hybrids between turgidum and timopheevii wheats are sterile and manifest a considerable amount of chromosomal irregularities in meiosis. On the basis of the available cytogenetic and molecular evidence, the genomic formula assigned to T. timopheevii (Zhuk.) Zhuk. is GGAA. In other words, GGAA T. timopheevii shares a similar genome A with BBAA T. turgidum and BBAADD T. aestivum, but does not contain their B genome (Sears 1969; Maan 1973). Moreover, even the genome A found in T. timopheevii is different from that encountered in T. turgidum and T. aestivum. It also contains a distinct cytoplasm (Maan 1973), which induces male sterility when combined with BBAA chromosomes. Domesticated Timopheev’s wheat is hulled, and shows close morphological similarities to domesticated emmer. It is an endemic crop restricted to western Georgia. The wild wheat from which it could have been derived is well known. The cultigens are fully inter-fertile and share identical chromosomal constitution with a group of brittle wild forms scattered over south-eastern Turkey, north Iraq, west Iran, and Transcaucasia. These were formerly named T. araraticum Jakubz. [=T. armeniacum (Jakubz.) Makush], but are now regarded as the

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wild race of the domesticated crop; i.e. T. timopheevii subsp. armeniacum (Jakubz.) van Slageren. (Please note that when we use the traditional classification, we refer to this wheat as T. araraticum). As already mentioned, armeniacum forms are genetically well isolated from both the wild and the domesticated forms of the tetraploid turgidum aggregate. However, morphologically they show striking similarities to wild emmer, T. turgidum subsp. dicoccoides. Moreover, in Turkey, Iran, and Iraq, both dicoccoides and armeniacum wheats are distributed over the same area (Map 4, p. 42) and it is practically impossible to separate their ripe ears from one another upon morphological examination (Tanaka and Ishii 1973). This can only be done by crossing and/or by molecular tests. For these reasons taxonomists dealing with the flora of south-west Asia (e.g. Bor 1968) frequently lump all Kurdish wild tetraploid wheat material together in what they call T. dicoccoides, disregarding the fact that they are actually confronted with two reproductively well-isolated entities. From the point of view of domestication and spread of domesticated wheats, the role of the tetraploid subsp. armeniacum GGAA stock is apparently negligible. With the exception of the much localized Georgian T. timopheevii cultivars, the chromosomes of the wild armeniacum stock are absent in the vast ensembles of cultivars and local land races of both tetraploid and hexaploid wheats. It could be argued that in the early days, both wild dicoccoides and wild armeniacum wheats could have been taken into cultivation in south-eastern Turkey, northern Iraq, and western Iran, and that the early non-brittle, hulled wheat remains from these places represent both stocks. Yet the question remains: if such alleged domesticated GGAA wheats were indeed produced in south-west Asia, why were they totally replaced by BBAA wheats, even among the local land races?

Barley: Hordeum vulgare Domesticated barley, Hordeum vulgare subsp. vulgare, is one of the main cereals of Mediterranean agriculture and a founder crop of Old World Neolithic food production. All over this vast area,

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barley is a universal companion of wheat, but in comparison with the latter it is regarded as an inferior staple and poor person’s bread. However, barley withstands drier conditions, poorer soils, and some salinity. Because of these qualities, it has been the principal grain produced in numerous areas, and an important element of the human diet. Indeed, barley has a much wider geographic distribution in the Mediterranean region and south-west Asia than wheat. Barley is also the main cereal used for beer fermentation. Preparation of this beverage seems to be a very old tradition (Hopf 1976; Darby et al. 1977; von Bothmer et al. 1991; Samuel 1996). The crop was, and still is, a most important feed supplement for domestic animals. Barley is an annual, predominantly self-pollinated diploid (2n = 2x = 14 chromosomes) grain crop. Consequently, its variation is structured in true breeding lines. Hundreds of modern varieties and thousands of land races are known. All cultivars have non-brittle ears; i.e. the spikes stay intact after ripening and are harvested and threshed by humans. This is in sharp contrast with wild barley forms, in which ears are always brittle. Non-brittleness in domesticated barley is governed by a mutation in either one of two tightly linked ‘brittle’ genes (Bt1, Bt2, Table 7, p. 61). The brittle wild-type allele in each locus is dominant; the non-brittle alleles are recessive. Like domesticated emmer, Btr1 and Btr2 are also located on the short arm of the homoeologous group 3 chromosome (3H). Many cultivars are homozygous recessive for both mutations. Others carry only one mutation (Takahashi 1964, 1972; Komatsuda and Mano 2002). Non-brittle mutations survive only under domestication, and non-shattering ears serve as a reliable indicator of domestication. Barley ears have a unique structure. They contain triplets of spikelets arranged alternately on the rachis (Hockett and Nilan 1985; Harlan 1995a). According to the morphology of the spikelets, barley under domestication can be divided into two principal morphological types: (i) Two-rowed forms—H. vulgare subsp. distichum, traditionally called Hordeum distichum L.—in which only the median spikelet in each triplet is both hermaphrodite and fertile, and usually armed with a prominent awn. The two lateral spikelets are males,

reduced, borne on longer stalks, and are grainless and awnless. Each ear thus contains two rows of grain-producing spikelets, one in each side (Fig. 15D). (ii) Six-rowed forms—H. vulgare subsp. Vulgare, traditionally referred to as H. hexastichum L.—in which the three spikelets in each triplet are hermaphrodite, bear grain, and usually all are awned. Ears in these varieties therefore have six rows of fertile spikelets, three in each side (Fig. 10E). The two-rowed condition is primitive; it is found in the wild progenitor of the crop as well as in all other Old World, wild Hordeum species. Sixrowed types of Hordeum vulgare were derived under domestication (Zohary 1969). A recessive mutation in the gene V and a dominant mutation in the gene i confer fertility to the lateral spikelets and cause the shift from two-rowed to six-rowed ears. The genotypic constitution of two-rowed forms is VVii. That of ordinary six-rowed cultivars is vvII (see Gymer 1978; Hockett and Nilan 1985). Wild barleys, as well as the majority of the domesticated forms have hulled grains; i.e. the pales are fused with the grains and cover them, even after threshing. In some domesticated varieties this hulled trait is lost. The ripe grains are free and are released, when the ears mature and dry, by threshing. In traditional farming communities, naked barleys were frequently favoured for the preparation of food, whereas hulled forms are preferred for brewing beer and for animal feed. The naked grain trait is controlled by a single recessive gene (n). Because of the striking differences in ear and grain morphology, taxonomists in the past frequently considered two-rowed barley forms and six-rowed barley forms as two separate species: H. distichum L. and H. hexastichum L. This nomenclature is still used by some botanists. Others divided the crop even further and called the forms containing fertile lateral spikelets on lax ears H. tetrastichum Körn., and those with dense ears H. hexastichum L. However, today we know (Zohary 1971; von Bothmer et al. 1991) that all domesticated barleys contain homologous chromosomes and are fully inter-fertile. Splitting is, therefore, genetically

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unjustified and the main domesticated barley types represent races of a single variable crop complex, H. vulgare L. (see Table 5). Carbonized grains are the most common barley remains found in archaeological excavations. Occasionally whole triplets are also retrieved, making it possible to determine whether the cereal was of the two-rowed or six-rowed type. Six-rowed barley forms can also be distinguished from two-rowed barleys by the morphology of the grains. In tworowed barleys all kernels are straight and symmetrical. In six-rowed barleys the lateral grains are often slightly bent and somewhat asymmetrical (Fig 16). The position and structure of the scars of the spikelets on the rachis internode provide an additional diagnostic trait to identify six-rowed barley in charred remains. The grains of naked barley (often called var. coeleste or var. nudum) are commonly recognized by their somewhat shriveled skin and by the furrow that stays narrow also near the apex (Figs 15 and 16). In summary, remains of two-rowed barley triplets in early archaeological sites can be very useful for the identification of the start of barley domestication. As with wheats, the disarticulation scars in the wild, brittle forms are smooth, whereas in non-brittle domestic varieties, threshing produces rough breakage scars. However, since in wild barley the first and second lowermost triplets (at the base of the ear) do not disarticulate easily, rare triplets with rough scars can be produced also upon reaping (and threshing) of wild spontaneum stands. As argued by Kislev (1997a), claims for barley domestication cannot be based on finds of rare breakage scars among numerous smooth ones. Only a prevalence of triplet remains with rough scars is a reliable indication of domestication. For the same reason, another good indication is finds of non-basal rachis fragment with more than one triplet.

Wild ancestry The wild ancestor of the domesticated barley is well known (Harlan and Zohary 1966; Zohary 1969). The crop shows close affinities to a group of wild and weedy barley forms which are traditionally grouped in Hordeum spontaneum C. Koch, but which are, in

53

fact, the wild race or subspecies of the domesticated crop. The correct name for this wild type is therefore H. vulgare L. subsp. spontaneum (C. Koch) Thell. (Table 5 and Fig. 15). These are annual, brittle, tworowed, diploid (2n = 2x = 14 chromosomes), predominantly self-pollinated barley forms, and the only wild Hordeum stock that is cross-compatible and fully inter-fertile with the domesticated barley. Vulgare x spontaneum hybrids show normal chromosome pairing in meiosis. Morphologically, the similarity between wild spontaneum and domesticated two-rowed distichum varieties is striking. They differ mainly in their modes of seed dispersal. Spontaneum ears are brittle and at maturity they disarticulate into individual arrow-like triplets. These are highly specialized devices which ensure the survival of the plant under wild conditions. Under cultivation this specialization broke down and non-brittle mutants were automatically selected for in the manmade system of reaping, threshing, and sowing. The close genetic affinities between the domesticated crop and wild spontaneum barleys are indicated also by spontaneous hybridization that occurs sporadically when wild and tame forms grow side by side. Some of these kinds of hybridization products, combining brittle ears and fertile lateral spikelets, were in the past erroneously regarded as genuinely wild types and even given a specific rank (H. agriocrithon Åberg). Extensive isozyme, seed storage proteins, and DNA tests have been carried out in barley (Nevo 1992). The results confirm the close relationships between the wild and domesticated entities grouped in the H. vulgare crop complex. They also clearly show that genetic polymorphism in the spontaneum wild populations are much wider than that present in the domesticated gene pool. Hordeum vulgare subsp. spontaneum is spread over the east-Mediterranean basin and the west Asiatic countries (Map 6), penetrating as far as Turkmenia, Afghanistan, Ladakh, and Tibet. Wild barley occupies both primary habitats and segetal, manmade, habitats. Its distribution centre lies in the Fertile Crescent, starting from Israel and Jordan in the south-west, stretching north towards south Turkey and bending south-east towards Iraqi Kurdistan and south-west Iran. In this area, wild spontaneum barley

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is continuously and widely distributed. It constitutes an important annual component of open herbaceous formations, and it is particularly common in the summer-dry deciduous oak park-forest belt, east, north, and west of the Syrian Desert and the Euphrates basin, and on the slopes facing the Jordan Rift Valley. From here, H. vulgare subsp. spontaneum spills over the drier steppes and semi-deserts. In south-west Asian countries, wild barley also occupies a whole array of secondary habitats; i.e. opened-up Mediterranean maquis, abandoned fields, and roadsides. It also infests cereal cultivation and fruit-tree plantations (Harlan and Zohary 1966). Further west, it is found in the Aegean region, the Mediterranean shore of Egypt and Cyrenaica, and extends further west to Algeria and Morocco. Further east, in north-east Iran, Central Asia, and Afghanistan, wild spontaneum barley is much more sporadic in its distribution; it rarely builds large stands and seems to be restricted, in most localities, to segetal habitats, ruins, or to sites which have been drastically churned by human activity. In general, wild barley does not tolerate extreme cold and it is only occasionally found above 1500 m above sea level. It is almost completely absent from the elevated continental plateaus of Turkey and Iran. On the other hand, it is somewhat more drought resistant than the wild wheats and penetrates relatively deeply into the warm steppes and deserts. The origins of domesticated barley are still not fully understood. Early crossing experiments and chloroplast DNA typing have suggested that domesticated barley evolved from one, two, or very few events (Zohary 1999). Later, Badr et al. (2000) examined 400 AFLP loci in 317 wild and 57 domesticated lines and found that barley is probably of a monophyletic origin resting in the Israel-Jordan area. However, in recent studies that included sequencing of seven genetic loci, Morell and Clegg (2007), and also Molina-Cano et al. (2005), Saisho and Purugganan (2007) and Wang and Ding (2009) suggested two origins: one within the Fertile Crescent and a second independent domestication farther east, possibly at the eastern edge of the Iranian Plateau. Apparently, the European and north African barley were largely connected to the Fertile Crescent while much of Asian barley is connected to the eastern center.

u A

B

Fig. 15 Continues overleaf

C

1

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55

F

G

D

H

E

Fig. 15 Main types of barley, Hordeum vulgare. A–Ear of wild barley, H. vulgare subsp. spontaneum [= H. spontaneum]. The ear is structured by seed-dispersal units (disarticulation units, spikelets) that disarticulate and shatters upon maturity. B–Seed-dispersal unit in dorsal side. C–Seeddispersal unit in ventral side, notice the upper (u) and lower (l) disarticulation scars. Note the ‘arrowhead-shape’ of the seed-dispersal units, with long and sturdy awns (partially removed in C). D–Ear of domesticated two-rowed barley, H. vulgare subsp. distichum. E–Ear of domesticated six-rowed barley, H. vulgare subsp. vulgare. F–Grain of hulled barley. G–Grain of naked barley. H–Triplet of six-rowed cultivated barley. Ears 1:1; grains 3:1 (Schiemann 1948).

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DOMESTICATION OF PLANTS IN THE OLD WORLD

A

0

5 mm

B

C

Fig. 16 Comparison between carbonized grains of wild and domesticated barleys. A–Wild barley, Hordeum vulgare subsp. spontaneum [= H. spontaneum] from final Mesolithic Mureybit, Syria (van Zeist and Casparie 1968). B–Grains of domesticated barley, H. vulgare subsp. vulgare. C–Grains of naked barley, H. vulgare var. nudum from Medieval Archsum, Germany (Kroll 1975). The arrow indicates lateral, twisted, grain; characteristic of the six-rowed barley cultivar.

Archaeological evidence Barley first appears in several pre-agriculture or incipient sites in south-west Asia. The remains are of brittle, two-rowed forms, morphologically identical with present-day wild spontaneum barley, and apparently collected from the wild (Fig. 16). The earliest records of such wild barley harvest comes from ca. 50,000 BP Kebara Cave (Lev et al. 2005) and from ca. 23,000 cal BP Ohalo II, a submerged Early Epi-Palaeolithic site on the south shore of the Sea of Galilee, Israel (Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004, 2008). In Ohalo II, the remains of H. spontaneum were found together with wild emmer wheat. Other early signs of H. spontaneum collection from the wild come from ca.15,500–10,150 cal BP, Franchthi Cave, Greece, from ca. 11,800–11,300 cal BP Mureybit (van Zeist and Casparie 1968; van Zeist and BakkerHeeres 1986), from ca. 10,500–10,200 cal BP Tell Aswad, east of Damascus, Syria (van Zeist and Bakker-Heeres 1985), and from ca. 11,700–10,550 cal BP Pre-Pottery Neolithic A, Netiv Hagdud, north of Jericho, Israel (Kislev 1997; Hartmann 2006; Weiss et al. 2006). An unusual find is a hoard

of approximately 260,000 grains (and 12,000 wild oat grains) from ca. 11,700–10,550 cal BP Gilgal, Israel, which most probably represents early, predomestication, farming (Hartmann 2006; Weiss et al. 2006). Wild barley is available from the ca. 9,600–8,750 cal BP Bus Mordeh phase of Ali Kosh, Iran (Helbaek 1969), the earliest layers, ca. 10,250– 9,550 cal BP, of Çayönü, Turkey (van Zeist 1972; van Zeist and de Roller 1991–2, 1995), and from ca. 8,700 BP pre-ceramic Beidha, Jordan (Helbaek 1966c). At the last three sites, brittle spontaneumtype barley was found in contexts showing definite signs of domesticated wheat. Remains of non-brittle two-rowed barley— domesticated forms that could survive only under cultivation—came from ca. 10,200–9,550 cal BP Middle PPNB (phase II) in Tell Aswad (van Zeist and Bakker-Heeres 1985), and from ca. 9,450–9,300 cal BP Neolithic Jarmo, Iraq (Helbaek 1959a, 1960; Braidwood 1960). In the latter site, Helbaek (1959b) was the first to show two-rowed barley remains still closely resembling wild spontaneum but also displaying a non-brittle rachis. Similar finds were reported by Hopf (1983) in ca. 9,900–9,550 cal BP pre-pottery Jericho. Indicative clues come from Ali

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57

Table 5 Taxonomy of the barley crop complex: Species according to traditional classification and their modern ranking on the basis of cytogenetic affinities Traditional classification

Modern grouping

Section Cerealia Åberg within the genus Hordeum L. containing the following species: 1. Wild two-rowed barley H. spontaneum C. Koch Brittle, hulled. 2. Domesticated two-rowed barley H. distichum L. Non-brittle, mostly hulled. 3. Domesticated six-rowed barley H. vulgare L. [= H. hexastichum L.] Non-brittle, both hulled and naked forms 4. Brittle six-rowed barley H. agriocrithon Åberg.

A single species containing both wild and domesticated forms. Collective name: H. vulgare L. 1. H. vulgare subsp. Spontaneum 2. H. vulgare subsp. distichum [= H. vulgare convar. distichon]

Kosh (Helbaek 1969), where the brittle spontaneumlike material characterized the lower layers, dated to ca. 9,600–8,750 cal BP, and in the upper strata dated to ca. 9,400–9,250 cal BP, it was replaced by non-brittle, broad-seeded, domestic forms. Hulled barley has been reported in ca. 10,650– 9,550 cal BP Early PPNB Kissonerga-Mylouthkia (Murray 2003) and Shillourokambos (Willcox 2000), Cyprus. Their domesticated status was based on grain size, while chaff remains are either wild-type or badly preserved. The domestic status of these finds requires further confirmation by chaff remains. However, as einkorn and emmer were found alongside with barley, and as no wild wheats grow in the island, it seems that cereals were introduced to the island from outside already at this early stage. Domesticated barley continues to be a principal grain crop in the south-west Asia throughout the Neolithic period. Its remains have been recovered side by side with wheats, in most Neolithic sites in which rich plant remains were retrieved. Shortly afterwards, we are faced with more advanced forms; i.e. six-rowed hulled as well as naked cultivars of barley. Remains of six-rowed barley start to appear in Turkey already in ca. 9,350–8,950 cal BP Aceramic Neolithic Çatalhöyük, while in the ca. 8,950–8,350 cal BP Ceramic Neolithic phase, it became firmly established (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007). In ca. 8,200–7,800 cal BP Late Neolithic Hacilar, it is represented by both hulled and naked varieties (Helbaek 1970). In the earliest phases of Ali Kosh,

Iran (Bus Mordeh and Ali Kosh, ca. 9,600–9,250 cal BP), only two-rowed barley occurs. But in the Mohammed Jaffar phase (ca. 8,400–8,350 cal BP), some six-rowed elements, as well as naked kernels, appear among the otherwise two-rowed material (Helbaek 1969). During the eighth millennium BP, hulled six-rowed barley establishes itself as the main cereal of the Mesopotamian basin as in ca. 7,300–7,000 cal BP Tell-es-Sawwan, Iraq (Helbaek 1964b). Hulled barleys are common in south-west Asia also during the Chalcolithic and Bronze Age. In these periods they show a tendency to outnumber the wheats. Being less sensitive to changes in climate and soils, barley adapted more easily to extreme conditions and probably replaced wheats on depleted soils, or in irrigated areas suffering from salinization. Barley was one of the principal crops that spread grain agriculture from south-west Asia, first to the Aegean region and subsequently to the Balkan countries, Caucasia, west Mediterranean basin, and central Europe, as well as west Europe, Egypt, and Transcaucasia (Maps 1 and 2). During the ninth millennium BP, barley emerges in the Aegean region and Greece as a constant companion of emmer and einkorn wheats. It is represented by both two-rowed and six-rowed forms and also by naked varieties (practically all Neolithic naked barleys are six-rowed). Such sites are Early Neolithic Franchthi Cave (Hansen 1991a, 1992), Aceramic Neolithic Cap Andreas-Kastros (van Zeist 1981) and Sesklo (Hopf 1962; Kroll 1981a), Greece, Aceramic Neolithic Dhali Agridhi, Cyprus (Stewart 1974), and Knossos, Crete (Sarpaki 2009). In Chalcolithic and

3. H. vulgare subsp. vulgare [= H. vulgare convar. vulgare] 4. Agriocrithon forms are now known to be secondary hybrid derivatives between 1 and 3.

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DOMESTICATION OF PLANTS IN THE OLD WORLD

0 100 200 miles 0

200

400 km

Map 6 Geographical distribution of wild barley, Hordeum vulgare subsp. spontaneum [= H. spontaneum]. The area in which wild barley is widely distributed is shaded. Dots represent additional sites, mainly of weedy forms. Wild barley extends eastwards beyond the boundaries of this map as far as Tibet and west China (Harlan and Zohary 1966).

Bronze Age Greece, barley is again gaining in importance and frequently becomes the prevailing cereal. At the same time, barley spread to Central Asia ca. 8,200–7,850 cal BP Jeitun, Turkmenistan, where the six-rowed form was found (Charles and Hillman 1992; Harris and Gosden 1996). Together with emmer and einkorn wheats, barleys emerge as one of the principal cereals of the Linearbandkeramik farmers that started Neolithic agriculture in central Europe in the first and second half of the eighth millennium BP. In eighth millennium BP, Neolithic cultures in the Balkans (notably in Bulgaria, see Marinova 2004, 2006), central and west Europe (e.g. Araus et al. 2007; Rottoli and Pessina 2007), one encounters mainly six-rowed forms and mostly naked grains from early on. Barley is relatively rare in the Linearbandkeramik culture. In some sites, naked grains are as common as hulled ones. In central Europe (as well as northern Europe), its importance clearly increases in later Neolithic times and especially in the Bronze Age.

In the Impressed Ware and later Neolithic cultures of the west Mediterranean basin (eighth through to the sixth millennia BP), barley remains are plentiful and they consist mainly of hulled and naked six-rowed forms (Hopf 1991). It remains a close associate of free-threshing wheats in these territories throughout the Bronze Age. Barley is the main companion of emmer wheat in the Neolithic settlement of the Nile Valley in the second half of the eighth millennium BP (Wetterstrom 1993) and also in this area it maintained its important role in food production through Neolithic and Bronze Age times. Barley also seems to be a main element in the diffusion of south-west Asian agriculture towards the east. It is present in the crop assemblage in several sites in Caucasia and Transcaucasia (Lisitsina 1984), in the first half of the eighth millennium BP in Caucasia, and in the second half in Transcaucasia. In conclusion, the archaeological finds show barley as a founder crop of the south-west Asian Neolithic agriculture and as a close companion of

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emmer and einkorn wheats. It is also clear that wild spontaneum barley is the ancestral stock from which domesticated barley was derived. This wild barley is a common annual plant in the Fertile Crescent and the area of its distribution largely coincides with the sites of the earliest finds of barley cultivation. This coincidence indicates conclusively that south-west Asia is the place of origin of domesticated barley. The archaeological remains enable us to trace the main developments of barley under domestication: first, the fixation of non-brittle mutations, and subsequently, the emergence of sixrowed hulled, and naked types. The principal role of barley in food production in the Old World in Neolithic and Bronze Age times is presently well documented.

Rye: Secale cereale Rye, Secale cereale subsp. cereale, is a characteristic grain crop of the temperate regions of the Old World. It is particularly appreciated in northern and eastern Europe because of its winter hardiness, resistance to drought, and its ability to grow on acid, sandy soils. Thus it succeeds under conditions in which wheat frequently fails (Evans 1995). Rye grains contain appreciable amounts of proteins and can be baked into dark-coloured ‘rye-bread’. Much of the present world production of rye is consumed in the form of bread, appreciated for its flavour and distinctive dense texture. The grains are also used as a high energy animal feed and for the preparation of rye whisky. The green plants are also commonly used for fodder. In contrast to most grain crops that are self-pollinating, rye is a crosspollinated cereal. Yields depend, among other factors, on effective wind pollination.

Wild ancestry Domesticated rye belongs to the small genus Secale L. Its diversity (in the wild) is centered in central and west Asia. Despite relatively large numbers of studies performed in this genus, its taxonomy is still somewhat uncertain. A widely accepted classification of this genus is the one proposed by Sencer and Hawkes 1980 (Zohary 1971; Stutz 1972; Kobylyanskyi 1989) and recently found to agree

59

with rDNA ITS sequence (de Bustos and Jouve 2002) and microsatellite markers (Shang et al. 2006). The following four biological species are now recognized. All are diploids (2n = 2x = 14 chromosomes) and are cold-tolerant. There is wide variation within this genus, both in the cultivars and in the wild forms—inside populations and between them. This variability is due to the fact that rye is largely cross-pollinated and a perennial plant. As a result of this variability, classification is a complicated task in this small genus. These are the current main four taxonomic groups in the genus Secale (Table 6): 1. The crop complex of S. cereale L. which contains the domesticated varieties, as well as con-specific weedy races and wild forms. All are annual, selfincompatible, chromosomally homologous, and fully inter-fertile with one another. Hybrids between the domesticated various forms are fully, or almost fully, inter-fertile with one another and show normal formation of seven bivalents in meiosis. In the past, rye taxonomists (such as Roshevitz 1947) split this complex into several species. The domesticated varieties were grouped in S. cereale L., while the variable weeds and wild types were treated as separate species (S. segetale (Zhuk.) Roshev., S. afghanicum (Vav.) Roshev., S. dighoricum (Vav.) Roshev., S. ancestrale Zhuk., and S. vavilovii Grossh.). Yet the results of comprehensive cytogenetic tests (Nürnberg-Krüger 1960a, 1960b; Khush 1963b; Stutz 1972; Vences et al. 1987) indicated that such splitting is unjustified. The species described are now regarded as main races or subspecies of the crop complex (Sencer and Hawkes 1980; Kobylyanskyi 1989; Shang et al. 2006; de Bustos and Jouve 2002). The variable S. cereale complex is currently roughly divided into the following principal races: (i) Domesticated varieties: These are non-shattering annual, self-incompatible plants with characteristic large and plump grains (Fig. 17C). (ii) Non-shattering weeds: This is a variable aggregate of obligatory weeds infesting wheat cultivation in Turkey, adjacent areas in Syria, Iraq, and Iran,

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DOMESTICATION OF PLANTS IN THE OLD WORLD

as well as the Balkan countries, Caucasia, and Transcaucasia. The mature ears of these weeds do not shatter and they mimic wheat in grain size and grain weight. Consequently, they are harvested and threshed together with the wheat. Since traditional winnowing does not separate grains of rye from those of wheat, rye seed is included in the harvest and planted with the wheat in the subsequent year. Farmers, particularly those in the elevated plateau of Anatolia and Armenia, tolerate some rye-weed infestation in their wheat crop, and for a good reason: in bad years with extreme cold and dry weather the rye weeds survive when wheat does not— ensuring the population a supply of what is sometimes referred to as the ‘wheat of Allah’ (Hillman 1978). (iii) Semi-shattering weeds: These weedy ryes are common in north-east Iran, Armenia, Afghanistan, and adjacent central Asian republics. Partially shattering ryes may infest wheat and barley cultivation, particularly irrigated fields. The rachis is semi-brittle; the upper part of the mature ear shatters spontaneously while the lower part stays intact and is reaped together with the wheat crop. Different populations vary in the degree of ear shattering. Frequently twothirds of the seeds produced undergoes ‘wildtype’ dissemination and are dispersed in the domesticated field. The remaining seeds are taken by the farmer to the threshing floor. (iv) Fully shattering wild types: In these forms the rachis is fully fragile and the mature ear disar-

ticulates spontaneously into individual spikelets. The kernels are narrow and fully covered by the brittle glumes. The individual wedgelike spikelets serve as seed-dispersal devices. The first brittle S. cereale forms were discovered near Aydin, western Turkey, by the Russian botanist P.M. Zhukovsky, and named by him ‘S. ancestrale’(Fig. 17B). Today, this robust rye type is known also from several other localities in the Izmir area. But since this rye infests fig plantations, roadsides, and vineyards, it is obviously a weed and not a genuinely wild type. Truly wild-rye forms, chromosomally homologous with the crop, occur in Armenia and in eastern Turkey (Map 7). They were first discovered in Armenia, where they have been named Secale vavilovii Grossh. Subsequently, large populations of these brittle, annual, cross-pollinated vavilovii forms were found also in eastern Turkey (Stutz 1972; Sencer and Hawkes 1980; D. Zohary unpublished data). This wild rye thrives on basaltic bedrocks. On the lower slopes of Mt Ararat, and in similar open terrain in adjacent volcanic areas, it frequently builds extensive stands. Compared to the weedy and robust ancestrale rye, vavilovii plants are somewhat shorter (usually 50–80 cm tall), but they grow in primary habitats and are obviously truly wild. Since they are fully inter-fertile with the domesticated rye (Kostoff 1937; Stutz 1972; Sencer and Hawkes 1980) they are regarded as a wild race (subspecies) of S. cereale.

Table 6 Characteristics of the four taxonomic groups in the genus Secale Species

Shattering

Wild/domesticated/ weed

Annual/perennial

Pollination system

S. cereale (i) Domesticated varieties (ii) Non-shattering weeds (iii) Semi-shattering weeds (iv) Fully shattering wild types S. montanum S. iranicum S. sylvestre

Non-shattering Non-shattering Semi-shattering Fully shattering Fully shattering Fully shattering Fully shattering

Domesticated Weed Weed Wild Wild Wild Wild

Annual Annual Annual Annual Perennial Annual Annual

Cross-pollinating Cross-pollinating Cross-pollinating Cross-pollinating Cross-pollinating self-pollinating self-pollinating

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Table 7 Recessive mutations which changed wild-type trait into domestic-type trait in south-west Asian founder crops. These mutations are responsible for the shift from wild type seed dispersal to human-dependent crops. There is no conclusive information yet regarding the situation in bitter vetch. Crop

Wild-type trait

Domestication trait

Einkorn wheat Emmer wheat Barley Lentil Pea Chickpea Bitter vetch Flax

Shattering ears Shattering ears Shattering ears Dehiscent pod Dehiscent pod Dehiscent pod Dehiscent pod Dehiscent capsule

Non-shattering ears Non-shattering ears Non-shattering ears Dehiscent pod Indehiscent pod Indehiscent pod Indehiscent pod Indehiscent capsule

2. Secale montanum Guss. [= S. strictum (Persl.) Persl.]. This rye species comprises a variable group of perennial, cross-pollinated forms, native to elevated plateaus and mountain systems in south-west Asia (especially Turkey), as well as the Caucasus, north Iran, the south Balkans, south Italy, Sicily, and Morocco (Sencer and Hawkes 1980). This species was apparently never domesticated. Perennial montanum forms have a tufted growth habit and pressed, shattering ears (Fig. 17A, Plate 7). Both morphologically and cytogenetically, perennial montanum forms are more distant from the crop. They differ from the domestic and wild forms of S. cereale by two chromosomal translocations (Stutz 1972; Sencer and Hawkes 1980). However, hybrids between annual S. cereale and perennial S. montanum are easy to make and they are only semi-sterile. Also, S. montanum is morphologically very variable and includes many eco-geographic races, such as: S. dalmaticum Vis., S. anatolicum Boiss, S. ciliatoglume (Boiss.) Grossh., and S. kupriyanowii Grossh. Over considerable areas on the elevated plateaus of Anatolia, S. montanum grows side by side with S. cereale, the first as a common grass in the nonarable steppe habitats, and the second as a weed in adjacent domesticated fields. Spontaneous hybridization between these two cross-pollinated grasses was found to be rather frequent in such contact places, particularly at the edges of cultiva-

Number of recessive mutations involved 1 2 2 1 1 1 2 1

Source

Love and Craig 1924 Nalam et al. 2006 Takahashi 1955; Zohary 1960 Ladizinsky 1979 Waines 1975 Kazan et al. 1993 Ladizinsky and van Oss 1984 Gill and Yermanos 1967; Diederichsen and Hammer 1995

tion (Stutz 1972; D. Zohary unpublished data). Introgressive hybridization often characterizes both species in these areas. Thus, in their main geographic centre, S. montanum and S. cereale are not reproductively fully isolated from one another and they still exchange genes. 3. Secale iranicum Kobyl. This is an insufficiently known annual wild rye. It is similar, in its general habit, to the wild and weedy brittle forms of S. cereale. Yet it is widely divergent from the crop chromosomally. It is also a self-pollinated species, has relatively small seeds, and it is isolated from the crop (and other Secale species) by strong hybrid sterility barriers. Until now, S. iranicum is known only from two collections made by H. Kuckuck in 1956 near Hamadan, Iran. He erroneously identified his plants as S. vavilovii and under that name sent seed samples to several centres of rye research (see Kuckuck 1973), causing confusion among researchers as to the nature of S. vavilovii. In fact, the genetic tests performed by Nürnberg-Krüger (1960a, 1960b), Kranz (1961), Khush (1963a, 1963b), Pérez de la Vega and Allard (1984), and Vences et al. (1987)—assumed to have been carried out on S. vavilovii—were actually performed on this totally different taxon. Only Kostoff (1937), Stutz (1972), and Sencer and Hawks (1980) used genuine S. vavilovii in their tests. It seems that Kuckuck’s collections represent an additional, distinct Secale species. Indeed it was described as such by Kobylyanskyi (1989).

62

0 0

DOMESTICATION OF PLANTS IN THE OLD WORLD

100 200

200 miles 400 km

Map 7 Geographical distribution of brittle wild rye, Secale cereale subsp. vavilovii [= S. vavilovii] (based on Stutz 1972; Sencer and Hawkes 1980).

4. Secale sylvestre Host. This is an annual, selfpollinating rye, with characteristic long awns, native to the Aralo-Caspian basin. Morphologically, S. sylvestre is well separated from the former three rye species. It also differs from S. cereale by three chromosomal translocations, and from S. montanum by a single one. It is largely inter-sterile with both these species.

Archaeological evidence Very few remains of Secale have been discovered in the Neolithic and Bronze Age settlements in southwest Asia. This is surprising since Turkey, Armenia, and Iran harbour a wealth of wild and weedy forms of S. cereale, and charred grains (Fig. 18) of this cereal can be easily identified. The earliest remains come from Epi-Palaeolithic sites in the Upper Euphrates valley in northern Syria. Numerous charred grains, later identified as a mixture of both wild rye and wild einkorn wheat (Fig. 10), were retrieved in ca. 11,800–11,300 cal BP

Tell Mureybit (van Zeist and Casparie 1968; van Zeist and Bakker-Heeres 1986; Willcox and Fornite 1999). The narrow shape of the kernels (Fig. 10) indicates that they represent wild forms. Similar narrow, wild-type rye grains (either of Secale cereale subsp. vavilovii or of S. montanum) were found in the ca. 12,700–11,100 cal BP Epi-Palaeolithic Tell Abu Hureyra (Hillman 1975, 2000a; Hillman et al. 1989, 2001). On the basis of grain morphology, Hillman et al. (2001) suggested these were domesticated form, a view that was later criticized, because of lack of chaff and problematic dating (e.g. Nesbitt 2002). Wild rye grains also discovered in two PPNA northern Syrian sites, ca. 11,500–11,000 cal BP, Jerf el Ahmar (Willcox 2002; Willcox et al. 2008, 2009), and ca. 10,700–10,400 cal BP Djade el Mughara (Willcox et al. 2008). Later, domesticated rye was found in ca. 9,450–8,450 cal BP PPNB Can Hasan III (Hillmam 1972, 1978) and in very small quantities at the nearby ca. 9,350–8,950 cal BP site of Aceramic Neolithic Çatalhöyük East, Turkey (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007). At Can Hasan III

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63

A Fig. 17 Ears and grains of wild and domesticated ryes. A–Perennial, brittle, Secale montanum. B–Annual, brittle (or semi-brittle), S. cereale subsp. ancestrale. C–Domesticated, non-brittle, S. cereale subsp. cereale, variety Petkus. Ears 1:1, grains 3:1 (Schiemann 1948).

relatively plump grains were discovered, together with some non-brittle rachis segments. As argued by Hillman (1978), these finds suggest that in Anatolia, rye had already entered cultivation in early Neolithic times, either as a non-brittle obligatory ‘ryeweed’ infesting wheat fields, or as a full-

fledged domesticated cereal crop. Yet, no additional rye remains have been discovered in other Neolithic south-west Asian sites. The next record comes from ca. 4,000 BP Bronze Age levels of Alaca Höyük in north-central Anatolia (Hillman 1978). Here a pure hoard of carbonized large grains of S. cereale was

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DOMESTICATION OF PLANTS IN THE OLD WORLD

B Fig. 17 Continued.

C

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discovered, indicating that at that time rye was grown as a crop in its own right. Outside the Fertile Crescent, early information on rye is fragmentary (for review see Behre 1992). The earliest convincing finds in Europe are from the following sites: (i) Sammardenchia and adjacent Early Neolithic sites (ca. 7,550–6,450 cal BP), northern Italy (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007). (ii) Middle Neolithic, Bükk culture (ca. 6,950–6,650 cal BP) Šarišské MichalˇanyFedelemka, Slovakia (Hajnalová 1993; Hajnalová and Hajnalová 2004). (iii) Several late Neolithic Funnel Beaker (TRB) culture sites in north, central, and south Poland (Giżbert 1960; Wasylikowa et al. 1991). Only a limited number of rye grains were found in the Polish sites compared to the abundance of wheat or barley grains. (iv) Few contemporary sites in Rumania, particularly Gumelnitza a culture (second half of the sixth millennium BP) Mǎgura Coneşti (Cârciumaru 1996) where a hoard of some 1000 charred kernels was discovered, suggesting that rye was grown as a crop, and ca. 6,650–6,350 cal BP Poduri (Cârciumaru and Monah 1985; Monah and Monah 2008) Further evidence on rye cultivation in Europe comes from several Bronze Age settlements in the Czech Republic and Slovakia where Tempír (1966, 1969) discovered rich remains of carbonized grains. Bronze Age records of rye are available also from Rumania (Cârciumaru 1996); Austria (KohlerSchneider 2001, 2003); and from Moldavia and the Ukraine (Wasylikowa et al. 1991). But in most cases they represent rye grains contaminating other cereals. Somewhat later, rye appears in Iron Age settlements in Germany (Hopf 1982, Table 1), Denmark (Helbaek 1954), Poland (Willerding 1970), and Crimea (Januševič 1978)—usually admixed with barley or wheat. Also in Hasanlu, Iran (Tosi 1975), S. cereale appears at the end of the fourth millennium BP and seems to have been a staple crop throughout the Iron Age. Rye was part of Roman grain agriculture and was grown in the cooler northern provinces. Carbonized rye grains have been retrieved from several Roman frontier sites along the Rhine and the Danube (Hillman 1978), as well as from the British Isles (Jessen and Helbaek 1944).

0

65

5 mm

Fig. 18 Carbonized grains of domesticated rye, Secale cereale, Medieval Archsum, Germany (Kroll 1975).

In summary, the available evidence from the living plants points to the annual, brittle rye [= Secale cereale subsp. vavilovii] as the wild ancestor of the domesticated crop. Eastern Turkey and adjacent Armenia seem to be the probable place of origin. The archaeological record supports this notion. The earliest sign of rye, associated with agriculture, comes from central Anatolia. Compared to wheat and barley, remains of rye in archaeological excavations are discouragingly few. Additional finds are necessary in order to establish the place and pattern of mode of origin, and mode of the spread of the rye crop. The available evidence also suggests that rye evolved first as a tolerated weed, and was only later picked up as a crop. The data also indicate that variation build-up in this cereal and the impressive evolvement of weedy ryes could have been consid-

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erably enhanced by introgressive hybridization with perennial Secale montanum—the common wild rye element distributed over the elevated, continental parts of Anatolia and adjacent areas in southwest Asia. The recent discovery of early Neolithic rye in Italy and Middle Neolithic in Slovakia supports the view that, despite earlier assumptions, rye arrived in Europe not through the Caucasus but rather via the Aegean Basin and the south Balkan.

Common oat: Avena sativa Avena sativa L., common oat, is a major cereal crop, in traditional Old World grain agriculture. It is a close companion of wheats and barley. The crop succeeds well in moist climates of temperate latitudes as well as in summer-dry Mediterranean conditions. In north-west Europe, oat frequently thrives better than wheat and is cultivated as a principal crop (Leggett and Thomas 1995). The nutritive value of oat grain is high; it contains about 15–16% protein and 8% fat. The crop serves as a staple in human diet and as a high energy supplement for farm animals. Three cytogenetically distinct stocks of Avena L. occur under domestication. Each had an independent origin (Zohary 1971). Yet, only one species, common oat, A. sativa (hexaploid, 2n = 6x = 42 chromosomes), established itself as a principal cereal. Two other oats (A. strigosa Schreb. and A. abyssinica Hochst.) are minor crops of negligible significance The main oat crop, A. sativa, is very variable. It comprises numerous contrasting domesticated varieties, as well as weedy races and truly wild forms. All are inter-fertile with one another and share the same hexaploid genomic constitution (Loskutov 2001). All are characterized by nonshattering panicles. According to their response to threshing, they are placed in three taxonomic ‘species’ that are actually only races of the same complex crop. Most cultivars are hulled; i.e. their grains remain invested by the pales. In some of these hulled forms (A. sativa in the narrow sense), threshing results in pressure breakage of the spikelet’s rachilla at the base of the upper floret. The rachilla segment remains attached to the lower floret. In

other sativa cultivars (traditionally named A. byzantina C. Koch), the rachilla segment breaks at its base and remains attached to the upper floret. The basal floret shows an abscission scar. In several hexaploid cultigens (frequently called A. nuda L.) the grains are free and threshing releases the naked kernels. Nakedness is a derived trait and occurs only under domestication.

Wild ancestry Avena L. is a Mediterranean genus comprising some twenty-five annual species. The majority of the oat species are diploid (2n = 2x = 14 chromosomes), seven are tetraploids (2n = 4x = 28 chromosomes), and six are hexaploid (2n = 6x = 42 chromosomes) (Leggett and Thomas 1995). The domesticated sativa oats (subsp. sativa) show tight genetic affinities and close morphological similarities to a group of wild and weedy oats, which are widely distributed over the Mediterranean basin, among them are A. sterilis L. (Fig. 19), A. ludoviciana Dur., A. occidentalis Dur., and A. fatua L. These wild types are also hexaploid and contain chromosomes homologous to those present in the cultivars (Loskutov 2008). Moreover, all wild and domesticated hexaploid oats are interfertile and occasionally cross in nature. Because of these close affinities, sterilis, occidentalis, and fatua oats are now recognized as the wild races of the domesticated crop and are placed within the A. sativa crop complex (Malzew 1930; Ladizinsky and Zohary 1971; de Wet 1981). The sativa complex is, however, chromosomally distinct and reproductively isolated from all other Avena species. Molecular studies have also confirmed this. A set of 413 AFLP bands tested in the 25 Avena species indicate that the sativa complex is a distinct group (Fu and Williams 2008), which is characterized as the only hexaploid member in this genus. Zhou et al (1999) examined several A. sterilis accessions from the Iran–Iraq–Turkey region and cultivated accessions, and found close association between them based on 248 polymorphic RAPD markers. Along with Jellen & Beard (2000), they divided the hexaploid group using the 7C-17 separation translocation and suggested at least two independent paths of domestication: one from A.

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sterilis (with the translocation) to A. sativa, and one from A. sterilis to A. byzantina (without it). Their conclusion was that A. sativa and byzantina are distinct races of the hexaploid biological species and was domesticated independently of one another. Just as in wheats and barley (see above), domestication brought about a breakdown of the original way of seed dispersal. Wild and weedy oats produce highly specialized drill-type diaspores, with characteristic kinked awns, in order to disseminate their seed and to insert them into the ground. They shed their seed immediately after maturation. As already noted, domesticated varieties have non-shedding panicles. Reduction of the awns and the evolution of relatively compact panicles, are additional conspicuous developments under domestication. Two distinct modes of seed dispersal operate in the wild members of the A. sativa complex, and serve as a diagnostic trait for their evaluation. Most widespread are oats in which the spikelet shows a single disarticulation point at the base of the lower floret. Consequently, the whole spikelet (minus the glumes), with two to three invested kernels, constitute a ‘drill-type’ dispersal unit (Plate 8). This fruiting type was traditionally named A. sterilis. In other forms (conventionally placed under A. fatua) the rachilla of the spikelet disarticulates at each node. The florets, each with a single seed, disperse individually. Sterilis-type oats widely colonize the Mediterranean basin from the Atlantic coast of Morocco and Portugal in the west to the Zagros Mountains in the east. In south-west Asia, they frequently grow together with wild wheats and barley. In addition to massive occupation of primary habitats, sterilis oats colonize abandoned cultivated ground all over the Mediterranean region aggressively, and they grow as a noxious weed in wheat and barley fields, orchards, and roadsides. Most conspicuous is the variation in the size of the spikelet, its hairiness, and colour. Forms with smaller spikelets are sometimes referred to as A. ludoviciana Dur., and those with larger spikelets and three to four fertile florets are called A. macrocarpa Mönch. Fatua oat varieties are distinctively weedy. They are widely distributed over the whole belt of Old

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World agriculture where they infest cereal fields and grow at the edges of cultivation. Only rarely do they occupy primary habitats. Fatua forms also thrive in colder, more continental climates. In elevated mountain cultivation and on the northern fringes of west Asiatic grain agriculture, fatua plants sometimes replace sterilis oats entirely.

Archaeological evidence The only sign of oat cultivation in south-west Asia came from an unusual find of some 12,000 Avena sterilis grains, no chaff, from ca. 11,700–10,550 cal BP PPNA Gilgal, Israel. This hoard also contains about 260,000 grains of wild barley, and probably represents pre-domestication cultivation as part of a barley field (Hartmann 2006; Weiss et al. 2006). Otherwise, there is no sign of oat cultivation or domestication in Neolithic or Bronze Age sites in south-west Asia and the Mediterranean basin. This, in spite of the fact that wild sterilis and fatua forms are massively distributed over these territories and even grow together with wild wheats and wild barley. The few oat remains retrieved from Neolithic Fertile Crescent and European sites seem to represent only shattering wild or weedy sterilis or fatua forms (Fig. 19). Definite indications of domestication; i.e. remains of non-shattering sativa or byzantina plants with their characteristic plump seed (Fig. 20) appear first in Europe, but only in the fourth and third millennia BP contexts (Willerding 1970, pp. 345–6; Villaret-von Rochow 1971). The earliest sites to report A. sativa finds, although rare, are ca. 7,600–7,400 cal BP, Sacarovca, Moldavia (Januševič 1984; Kuzminova et al. 1998; Monah 2007b,), and ca. 6,650–6,350 cal BP Poduri, Rumania (Cârciumaru and Monah 1985; Monah and Monah 2008). Furthermore, Füzes (1990, as reported by Gyulai 2007) identified impressions with some grains of naked oat (Avena cf. nuda) from ca. 7,000–6,300 BP Aszód-Papi Földek, Hungry. Later, domesticated oats are present in fourth millennium BP Nitriansky Hrádok, Czech Republic (Tempír 1969; Kühn 1981), Slovakia (Tempír 1966), and ca. 2,900–2,800 cal BP Forsandmoen and adjacent sites in Norway (Bakkevig 1982, 1995).

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A

B

0

2 cm

Fig. 19 Wild oat, Avena sativa subsp. sterilis [= A. sterilis]. A–Fruiting panicle. B–Seed-dispersal unit (spikelet without glumes) (Malzew 1930, plates 86, 88).

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Broomcorn millet: Panicum miliaceum

0

5 mm

Fig. 20 Carbonized grains of domesticated oat, Avena sativa, Medieval Archsum, Germany Kroll 1975 (Kroll 1975).

Domestic oat becomes much abundant in central and western European contexts from the Roman Period onward. In conclusion, the data available from the archaeological excavations, as well as the evidence obtained on the ecology and distribution of the wild relatives, support the notion that domesticated A. sativa should be regarded as a secondary crop. Probably, oat started its evolution under domestication not as a crop but by evolving weedy types that infested wheat and barley cultivation. Only later were such weeds picked up and planted intentionally. In the temperate periphery of Old World agriculture, oats not only supplemented wheat and barley, but also evolved into a principal grain crop.

Broomcorn millet (common millet, Proso millet), Panicum miliaceum L. (Plate 9), ranks among the hardiest cereals. It is a warm-season crop which stands up well to intense heat, poor soils, and severe droughts, completing its life cycle in a very short time (60–90 days) and succeeding in areas with short rainy seasons. This is the true millet of classic times (the Romans’ milium and the Hebrews’ dokhan). Today, P. miliaceum is grown mainly in eastern and central Asia, in India and (sparsely) in south-west Asia. The de-husked grains are boiled and cooked like rice, or ground for the preparation of porridge. They are quite rich (10–11%) in proteins. The seeds are also used as birdfeed. Broomcorn millet is tetraploid (2n = 4x = 36 chromosomes). It is mainly self-pollinated. The wild ancestor of the domesticated P. miliaceum has not been satisfactorily identified. Weedy forms of this millet are widespread in central Asia, from the Aralo-Caspian basin in the west to Sinkiang and Mongolia in the East. They are referred to as P. miliaceum subsp. ruderale (Kitag.) Tzvelev [= P. spontaneum Lyssov ex Zhuk.]. Recently these weeds, with their characteristic shattering panicles, also spread to central Europe and north America (Scholz 1983). Probably, the vast semi-dry areas in central Asia harbour not only weedy, but also genuinely wild miliaceum forms. Lu et al (2009) have identified phytoliths and biomolecular components of certain P. miliaceum types dated to the eleventh to ninth millennia BP. However, there are still large complexities and points of debate regarding the crop/weedy type status of any findings of P. miliaceum (Hunt et al. 2008). Additional archaeobotanical finds and taxonomical studies are necessary in order to establish the origin and dispersal patterns of this crop.

Archaeological evidence Identifying Panicum miliaceum remains and differentiating it from those of Setaria italica can be problematic. Hunt et al. (2008), compiled a comprehensive review of early, pre-7,000 cal BP, finds of both Panicum and Setaria finds from archaeological sites across Eurasia.

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They also referred to issues of taphonomy and identification criteria offered so far, as well as the differentiation between wild/weed/crop status (see there). The earliest archaeobotanical P. miliaceum grains are reported from a number of early Neolithic sites in north China, e.g. 7,670–7,610 cal BP Xinglongwa (Zhao 2011) and ca. 8,060–7,750 cal BP, Yuezhuang site (Crawford et al. 2006). No early finds are available from central Asia. However, remains of millet have been reported from several Neolithic sites dated to the first half of the eighth millennium BP, such as ca. 8,000–7,150 cal BP Arukhlo 1 and Arukhlo 2, Georgia (Januševič 1984; Lisitsina 1984; Schultze-Motel 1988a), ca. 7,600–7,400 cal BP Sacarovca, Moldavia (Januševič 1984; Monah 2007b; Kuzminova et al. 1998), and ca. 7,650–7,400 cal BP Mohelnice, Czech Republic (Opravil 1979, 1981; Kühn 1981). Broomcorn millet is also reported from seventh millennium BP VI strata in Tepe Yahya, Iran (Costantini and Costantini-Biasini 1985). As Hunt et al. (2008) indicated, it is too early to determine whether these were locally domesticated or represent early east–west movement of a domesticated crop. However, the Georgian finds come from sites situated not far away from the geographic belt where P. miliaceum grows wild. Finds of the characteristically small oval seeds (Fig. 21A) of P. miliaceum continue to appear in late eighth and from the seventh millennia BP settlements in east and central Europe. The nature of its appearance is geographically sporadic throughout the Linearbandkeramik and later sites, but became more common in late Neolithic and in Bronze Age cultures. Some of these finds are ca. 7,250–6,650 cal BP Zánka-Vasútállomás, Hungary (Füzes 1990, 1991), ca. 6,500–6,000 cal BP Rivne, Ukraine (Pashkevich 2003), ca. 6,150 BP Domica (= Kesevo) in Czechoslovakia (Fietz 1936; Tempír 1969), and in seventh millennium BP Sesklo, Greece (Hopf 1962; Kroll 1981b). Van Zeist (1980) suggested that P. miliaceum reached the Mediterranean basin from the north or the north-east. The find from ca. 6,300–4,600 cal BP Monte Còvolo, Italy (Pals and Voorrips 1979), indicates that this was a rather early movement. Remains of P. miliaceum also occur in Tripolye culture (ca. 6,000–4,750 cal BP) sites such as Soroki in the Ukraine (Januševič 1976, 1978), in ca. 5,500 BP

A

B 0

5 mm

Fig. 21 Carbonized grains of: A–Broomcorn millet, Panicum miliaceum; B–Foxtail millet, Setaria italica. Bronze Age Kastanas, Greece (Kroll 1983).

Gumelnitsa culture contexts Morteni, Rumania (Cârciumaru 1996, p. 91), and in ca. 5,900–5,700 cal BP Gomolava, Macedonia (van Zeist 1975, 2003). Somewhat later, lumps of pure charred grains were retrieved from the Funnel Beaker (TRB) strata in Szlachcin, Poland (Wasylikowa et al. 1991). The earliest evidence for broomcorn millet in Austria came from ca. 5,050–4,750 cal BP late Neolithic Anzingerberg/Hundssteig (Kohler-Schneider 2007; Kohler-Schneider and Caneppele 2009). At about the same time, broomcorn millet arrived to ca. 4,800–4,200 cal BP Crasto de Palheiros, Portugal (Pinto da Silva 1976). During the Bronze Age, it appears in ca. 1,850–800 cal BC Zürich, Switzerland (Jacomet 1988, 2004;

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Jacomet et al. 1989; Brombacher and Jacomet 1997; Favre 2002; Brombacher et al. 2005), and to ca. 905– 869 BC Grésine (Bouby and Billaud 2001), and later in Ouroux-Marnay (Hopf 1985), France. In the north, the earliest finds include ca. 1,400–950 cal BC Lindebjerg and Voldtofte, Denmark (RowleyConwy 1979, 1983), and ca. 1,150–950 cal BC Borge Vestre, Norway (Sandvik 2007, 2008). This crop was also discovered from ca. 2,350–2,100 cal BC Harappan Shortughai, Afghanistan (Willcox 1991), and a large quantity of charred grains was uncovered in ca. 1,900–1,550 BC Bronze Age Haftavan, Iran (Nesbitt and Summers 1988). The first known deposit of identifiable grains in Iraq comes from ca. 700 BC Nimrud (Helbaek 1966b). Signs of the cultivation of P. miliaceum in the Levant appear relatively late – Middle Bronze Age Tell Mozan, Syria (Riehl 2000), and ca. 1,200–500 BC Iron Age Deir Alla, Jordan (Neef 1989). In conclusion, the evidence from the living plants and the archaeological remains is still fragmentary, but it does show that broomcorn millet is a relatively old crop, and that it does not belong to the south-west Asian Neolithic crop assemblage. It may be that P. miliaceum is an eastern or central Asian element that was picked up and added to wheats and barley agriculture soon after its arrival and establishment in these areas. The scanty data available do not rule out the possibility that P. miliaceum (together with foxtail millet, below) represents an independent experiment in domestication, both in East Asia and Caucasian-European region.

Foxtail millet: Setaria italica Foxtail or Italian millet, Setaria italica (L.) P. Beauv. (Plate 9), is a principle founder crop of Chinese grain agriculture. Genetic evidence suggests it was domesticated independently in other parts of Asia (Fukunaga et al. 2005). Italian millet and broomcorn millet are the major crops in Chinese dry-land agriculture of a wide loess area along the Yellow River, from southern Mongolian Steppe in the north and the Huai River in the south (Zhao 2011). It is widely cultivated today in India, China, and Japan, while it is much rarer in European and south-west Asian agriculture. This is a diploid (2n = 2x = 18 chromo-

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somes), predominantly self-pollinated millet with characteristic dense, bristle-bearing panicles, and small, oval grains which are tightly enclosed by their pales (Prasada Rao et al. 1987; de Wet 1995). Similar to the broomcorn millet (p. 69), Foxtail millet is a warm-season cereal. It survives well under dry conditions and completes its growth cycle in a relatively short time. The wild progenitor of foxtail millet is well identified. Domesticated S. italica shows close morphological affinities to, and is inter-fertile with, wild S. viridis (L.) P. Beauv., a common variable summer weed widely spread across Eurasia (de Wet et al. 1979). Molecular markers and components have revealed that the origin of foxtail millet domestication was probably in China (Le Thierry d’Ennequin et al. 2000; Lu et al. 2009) with the possibility of a few additional domestication events elsewhere in Eurasia (Fukunaga et al. 2005). The spontaneous and the domesticated foxtail millets differ from one another mainly in their seed-dispersal biology. Wild and weedy forms shatter their seed while the cultivars retain them. Genetic tests indicate that the shift is governed by two complementary recessive mutations (Prasada Rao et al. 1987). Under domestication, foxtail millet underwent changes in plant habit, including a reduction of the number of flowering tillers, and enlargement of the flowering panicles (de Wet 1995).

Archaeological evidence Setaria italica is the principal grain crop of the Neolithic agriculture in north China (Ho 1977; Zhimin 1989; Crawford 1992; Zhao 2011). Identifying Setaria italica remains, and differentiating it from those of Panicum miliaceum, can be problematic. Hunt et al. (2008) compiled a comprehensive review of early, pre-7,000 cal BP finds of both Panicum and Setaria from archaeological sites across Eurasia. They also referred to issues of taphonomy and identification criteria offered so far, as well as the differentiation between wild/ weed/crop statuses. The earliest archaeobotanical Setarica italica grains are reported from a number of early Neolithic sites in north China, e.g. 7,670– 7,610 cal BP Xinglongwa (Zhao 2011) and ca. 8,060–7,750 cal BP Yuezhuang site (Crawford et al.

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2006). Zaho (2011) stresses that broomcorn millet is always more abundant than foxtail millet in these early sites (which is true for later sites and regions as well). However, as indicated by Hunt et al. (2008), unlike broomcorn millet which occurs in pre-7,000 cal BC sites both in western Asia and in Europe, foxtail millet occurs, at this time, only in northern China. Soon afterwards, S. italica emerges as the principal cereal of the Yang-shao culture (seventh and fifth millennia BP). The finds demonstrate that in Yang-shao times, foxtail millet was occasionally accompanied by broomcorn millet, P. miliaceum (p. 69). Both millets continue to play a major role in north China’s food production today (Ho 1977). In Europe, carbonized seeds (which were definitely assigned to S. italica) first appear in the second millennium BC Bronze Age settlements in central Europe, like ca. 1,850–800 cal BC Zürich, Switzerland (Netolitzky 1914; Jacomet 1988, 2004, Jacomet et al. 1989, Brombacher and Jacomet 1997; Favre 2002; Brombacher et al. 2005) and in ca. 905– 869 cal BC Grésine, France (Bouby and Billaud 2001). This millet is also reported from Late Bronze Age Kastanas, Macedonia, Greece (Kroll 1983; see Fig. 21). Its earliest evidence in Austria came from Late Bronze Age ca. 1,200–700 cal BC Stillfried (Kohler-Schneider 2001, 2003). Additional finds are available from Iron Age Europe. The earliest definite evidence for foxtail millet cultivation in southwest Asia comes from Iron Age (ca. 600 BC) Tille Höyük, south-east Turkey (Nesbitt and Summers 1988). Here a large quantity of pure grains was uncovered, indicating the use of foxtail millet as a crop. The available archaeological evidence indicates that foxtail millet is a relatively old domesticant. It was probably first taken into cultivation in northeast Asia. Together with rice and common millet, they served as founder crops in the development of agriculture in north China. Like common millet, S. italica does not belong to the Neolithic south-west Asian crop assemblage. Moreover, it appears in Europe rather late. It is still an open question whether S. italica was taken into cultivation only in north China or also elsewhere in this vast area.

Latecomers: sorghum and rice Two additional principal cereals—sorghum, Sorghum bicolor (L.) Moench, and rice, Oryza sativa L.—are not indigenous to south-west Asia and the Mediterranean basin. They are latecomers that probably arrived in these areas as fully developed crops only in Greek and Roman times, or even later.

Sorghum Sorghum bicolor (L.) Moench is a warm-weather grain crop grown extensively in Africa, south-west Asia, and the Indian subcontinent. Both sorghum and pearl millet are able to tolerate very arid conditions and have a major role in food production is these areas. Wild forms closely related to the variable sorghum crop are confined to Africa south of the Sahara (including Yemen). This indicates that this grain crop must have been domesticated in this area, probably in the Savanna belt south of the Sahara (de Wet et al. 1976; Harlan 1992b; Wetterstrom 1998). Sorghum is a diploid (2n = 2x = 2 chromosomes), predominantly self-pollinated crop. During domestication, it underwent considerable differentiation. Five principal domesticated races have been recognized in this crop (Harlan and Stemler 1976). The wild relatives of the crop are very variable. Four principal eco-geographical wild races have been recognized in S. bicolor (de Wet et al. 1976, Map 1), as well as segetal forms (including companion weeds) that frequently infest sorghum cultivation. The wild and weedy forms are also diploid and predominantly self-pollinated. All are inter-fertile with one another as well as with the cultivars. Using cytological techniques and molecular markers such as mitochondrial ITS variation across all twentyfive species of Sorghum, the crop and its wild progenitors were grouped into a distinct phylogenetic lineage of Sorghum (Dillon et al. 2001; Price et al. 2005). Similar to rice (pp. 73–74), wild and weedy S. bicolor frequently hybridize with their domesticated counterparts. For this reason, in Africa, one is confronted with a variable crop complex comprising: (i) wild forms occupying more stable habitats; (ii) cultivars grown by the farmers; and (iii) weeds colonizing secondary habitats and infesting sorghum cultivation. The finds in pre-agriculture (ca 8,000

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BP) Nabta Playa, south Egypt, (Wasylikowa and Kubiak-Martens 1995) have shown that seeds of wild S. bicolor were collected from the wild before its domestication. The archaeological exploration of sub-saharan Africa is in its early stages, and we lack critical information for determining when and where sorghum could have been taken into cultivation. So far, remains of African domesticated sorghum are available only from late contexts. The richest finds, preserved by desiccation, came from Qasr Ibrim, Egyptian Nubia (Rowley-Conwy 1991; RowlyConwy et al. 1999; Clapham and Rowley-Conwy 2007). Remains of local wild forms such as S. bicolor (L.) Moench subsp. arundinaceum (Desf.) de Wet & Harlan, occur in contexts dated 800–600 BC, while domesticated sorghum, conforming in its morphology to race bicolor cultivars, abound in contexts dated from 100 AD to 1,200 AD. A bouquet of ripe panicles, dated 42–640 AD, intermediate in its morphology between bicolor and durra cultivars, was also retrieved, and by 1,200 AD the more advanced, free-threshing durra forms make their appearance. The earliest archaeological evidence on sorghum cultivation, available up to date, does not come from Africa but from the Indian subcontinent. Reports about the presence of sorghum are available from about a dozen of second millennium BC sites in Pakistan and India (see review by Fuller 2002). As Fuller remarks, in some of these cases, the identification is problematic. However, the remains uncovered from several other locations definitely belong to this crop. Since Sorghum bicolor does not grow wild in India, these finds have been interpreted (see Vishnu-Mittre and Savithri 1982; Harlan 1992b) as indicating: (i) an even earlier domestication in Africa, and (ii) an early migration of domestic sorghum, from East Africa into the Indian subcontinent. This interpretation got further support from the fact that several other African grain crops, namely: pearl millet Pennisetum glaucum (L.) R. Br., cow pea Vigna unguiculata (L.) Walp., and hyacinth bean Lablab purpureus (L.) Sweet, show similar patterns of migration. Their wild progenitors are restricted to Africa. They emerge (as crops) in the Indian subcontinent already in the second millennium BC. Further, this view of early spread

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was supported by the recent finds of ca. 2,500 BC domesticated pearl millet in Mali, west Africa (Manning et al. 2011). Sorghum seems to have arrived late in south-west Asia and the Mediterranean basin, even after the medieval period, in light of its absence from medieval sites in Syria (Samuel 2001: 429). As late as Roman times, this cereal was almost unknown in the belt of Mediterranean agriculture. Although reported (as a crop) from Qasr Ibrim, from 100 AD onwards (Rowley-Conwy 1991), sorghum cultivation seems to have stopped in Nubia and did not establish itself in Lower Egypt. The advanced durra cultivars which now characterize traditional summer cropping in south-west Asia, in India and in Pakistan, are not widespread in Africa. There, they are largely confined to the fringes of the Sahara and to Ethiopia. For these reasons Harlan and Stemler (1976) speculated that the more advanced, freethreshing durra forms evolved in India from more primitive bicolor material brought from east Africa, and that durra type sorghum was introduced to south-west Asia and north-east Africa rather late. However, this is just an attractive speculation. As argued by Rowley-Conwy et al. (1997, 1999), a sounder appreciation of sorghum domestication would be possible only when its start in Africa will be better understood.

Rice Oryza sativa L. is an east and south Asian element. It is the domesticated founder crop in this part of the World (Chang 1995) and is one of the major food suppliers to the human race. Rice is a diploid (2n = 4x = 24 chromosomes; genomic constitution AA), predominately self-pollinated. This is a very productive crop when grown in paddies, where fixed nitrogen is naturally provided by co-habiting blue-green algae. In the course of its long and complicated history of cultivation, Asian rice underwent considerable differentiation. Thousands of cultivars evolved, fitting the wide range of environmental conditions into which this crop complex has been introduced. They fall into two main groups: (i) short, thick grained ‘japonica’ (= ‘sinica’) forms, adapted to the relatively cool climate in northern

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China and south-east Asia; and (ii) long, thin grained ‘indica’ adapted to hot, tropical climates. However, numerous other varieties and subvarieties are recognized (see Fuller et al. 2010 for a detail treatment and updated archaeobotanical and genetic knowledge). The crop is closely related to, and fully inter-fertile with, a variable assemblage of wild and weedy rice forms which are widely distributed over south and east Asia including the Indian subcontinent, south China, and Indonesia (Chang 1995; Vaughan et al. 2008). They include both perennial and annual wild forms (frequently called O. rufipogon Griff. and O. nivara Sharma and Shastry respectively), as well as shattering weedy rices (known as spontanea forms of O. sativa), which frequently infest rice cultivation. In many places over these vast territories, the domesticated cultivars, the weedy forms, and the wild races grow side by side, hybridize with one another, and form a huge crop complex. There is an ongoing debate as to whether perennial O. rufipogon or annual O. nivara, or both, were the direct ancestors of O. sativa (Sweeney and McCouch 2007). The morphological and genetic separation between and within these taxa is, therefore, problematic. Recent genetic studies (Fuller and Sato 2008; Fuller et al. 2010) suggest that japonica rice was domesticated in the Yangtze River areas in China sometime about 10,000 years ago, and then spread westward, hybridized with the wild progenitor of indica rice, and this hybrid eventually domesticated as the indica rice somewhat around 4,000 BP. Since the distributional range of the wild relatives of domesticated Oryza sativa is immense, one can provide only a general orientation as to where the wild progenitor of rice was introduced into domestication. However, archaeological finds in China, India, and several countries in south-east Asia provide some clues as to where and when rice domestication originated (Sweeney and McCouch 2007; Fuller et al. 2010; Hosoya et al. 2010; Nesbitt et al. 2010).

Based on increases in domesticated-type spikelet bases, in rice grains, and in arable weeds typical of rice paddies, between strata, Fuller et al. (2009) suggested that rice’s pre-domestication cultivation began around 7,000 cal BP in the Lower Yangtze region, China, pre-dating its local domestication in around 6,000 cal BP. Somewhat later (in the seventh millennium BP), remains of domesticated rice appear at Chengtoushan in the middle Yangtze Valley (Fuller et al. 2007). Domesticated rice appears in central China in about 5,000–4,000 BP (Crawford and Shen 1998). The earliest convincing signs of rice cultivation in the Indian subcontinent come from contexts dated to the second millennium BC; when rice apparently became an important summer crop in the subcontinent (Nesbitt et al. 2010).The number of finds increases considerably from the second millennium BC onwards (Glover and Higham 1996; Meadow 1996; Fuller 2002). Towards the end of the third millennium BC, imprints of rice stalks and husks, as well as charred grains, appear in farming sites in the central and eastern parts of the Ganga Valley. By about 2,000–1,800 BC, rice seems to have been added effectively as a summer crop, to wheat and barley agriculture in the Indus Valley and adjacent areas, marking the beginning of the distinctive, two crop system of the Harappan agriculture. Asiatic rice was probably introduced into south-west Asia in Hellenistic times. Both Greek and Roman writers knew about this cereal and said that it was grown in Baktria, Mesopotamia, and Syria (Lenz 1859, pp. 229–30). A large sample of rice grains was retrieved from an AD first-century Parthian grave, Ville Royale II, Susa, Iran (Miller 1981). In Roman times, highly prized large kernel rice was produced in Israel (Feliks 1983) and imported rice is known from ca. 50 BC Berenike on the Red Sea coat of Egypt (Cappers 2006). At this time, rice was also grown in the Po Valley in Italy.

C H A PTER 4

Pulses

Annual legumes (from the Papilionaceae/Fabaceae family, of the Leguminosae) cultivated for their seed accompany the cereals in most regions of grain agriculture. They are attractive because contrary to most other flowering plants, legumes are able to fix atmospheric nitrogen through symbiosis with the root bacterium Rhizobium. Rather than use nitrogen up, pulses add it to the soil. By practising rotation or mixing of legume crops with cereals, the cultivator is able to maintain higher levels of soil fertility. Another virtue is that the seeds of pulses are exceptionally rich in storage proteins, whereas grass kernels are rich in starch. Therefore they complement each other as food elements and contribute to a balanced human diet. In traditional agricultural communities, pulses served—and still serve—as a main meat substitute. Both the agronomic compensation and the dietary complementation between cereals and pulses were appreciated in the early days of agriculture. Each major agricultural civilisation developed not only its staple cereals, but also its characteristic companion legumes. Wheats and barley agriculture in west Asia and Europe had pea, lentil, faba bean, and chickpea. Maize in Meso-America was accompanied by several species of Phaseolus beans, and in South America also by the groundnut. Pearl millet and sorghum cultivation in the African Savannah belt were associated with cowpea and Bombara groundnut. Soybean was added to cereal cultivation in China. Hyacinth bean, black gram, and green gram were introduced into agriculture in India. Pulses seem to have started their role as companions of wheats and barley very early in the agricultural history of the Old World. The available archaeological evidence indicates that pea, lentil,

chickpea, and bitter vetch (probably also grass pea) were introduced into cultivation more or less together with the principal cereals. Their remains abound in south-west Asian Neolithic settlements. Some of them (particularly lentil and pea) are common in the contexts of the Neolithic sites that appeared soon afterwards all over the vast area from the Atlantic coast of Europe to the Indian subcontinent. In other words, in south-west Asia, wheats and barley were not domesticated alone; pulses accompanied them from the very start (Zohary and Hopf 1973; van Zeist 1980; Smartt 1990; Zohary 1996; Butler 1998 Butler 2009). Several other pulses became grain crops in west Asia and in Europe only after the establishment of the ‘first wave’ of legume crops. Prominent among them were the faba bean and fenugreek. They were followed, apparently later, by the white lupin. The cowpea, Vigna unguiculata, was introduced into cultivation in Africa south of the Sahara and reached the Mediterranean basin only in classical times. Self-pollination seems to have been a major asset in domestication also in the seed legumes. Similar to the cereals (pp. 20–23), the majority of Neolithic ‘first wave’ domesticates (pea, lentil, chickpea, and bitter vetch) are predominantly self-pollinated. So are their wild progenitors. Thus, in this group of grain crops, wild self-pollinated candidates were ‘pre-adapted’ for domestication as compared with cross-pollinated plants. The advantages conferred by self-pollination are the establishment of a barrier between wild and cultivated populations, and the automatic fixation of desired genotypes. As in the cereals (pp. 22–23), the evolution under domestication of the seed legumes is characterized by the development of a syndrome of

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domestication traits (Harlan 1975; Hammer 1984; Smartt and Hymowitz 1985; Harlan 1992a). Very likely, many of these domestic traits evolved as a result of predominant self-pollination and unconscious selection (Zohary 1989b). First, in the wild type, there is an automatic selection for the breakdown of seed dispersal and the selection for the retention of seeds in the pod. In most pulses (e.g. pea, lentil, the various members of the genus Vicia) seed dispersal in the wild depends on the bursting of the mature pods. The pods of domesticated forms do not burst or at least do not split open quickly. They are harvested and threshed by the farmer. In pea, lentil, chickpea, and bitter vetch, the change to non-dehiscing or slow-splitting pods is a simple genetic event. It is controlled by a single recessive mutation (or two such mutations, Table 7, p. 61). A second change is the loss of the wild-type seed dormancy. The seeds of most wild Mediterranean pulses have a relatively thick and coarse seed coat, which insulates the seeds from water penetration, and spreads germination over several years. This wild-type adaptation breaks down under domestication. Seeds of the domestic varieties tend to have thinner seed coats and are permeable to water. Consequently, in the traditional Mediterranean pulse crops, practically all the fresh seeds germinate the same year they are sown. In other words, the introduction of the wild progenitors into a system of reaping and sowing brought about an automatic selection of mutations causing a breakdown of the wild-type germination inhibition. At least in some pulses, the seed coat structure can be used—as a telltale trait—to identify domestication. However, this is only occasionally feasible in archaeobotanical studies. Most of the time, charring pulverizes the seed coat, leaving only the cotyledons and the embryo intact. Seed coats persist only in exceptional cases of preservation. Another trend under domestication is the change in seed dimensions (Fig. 22). Most pulse crops bear considerably larger seeds than their wild ancestors. In some cases, there is a three- or fourfold increase in the volume and the weight of the seed (Zohary and Hopf 1973; Smartt 1990). This increase has been a gradual process. Seeds retrieved from Neolithic contexts are still relatively small and not very different in size from those of their wild relatives. The large-seeded pulses with which we are familiar

today reached their present large dimensions only in classical times. In several legumes, domestication brought about striking changes also in the plant habit. The wild relatives of the cultivated pea, grass pea, and the

A

B

C

D

E

Fig. 22 Archaeological carbonized seed remains (left) and, for comparison, seeds from modern cultivars (right) of the first five south-west Asian domesticated pulses. A–Pea, Pisum sativum, carbonized seeds from Early Bronze Age Arad, Israel. B–Lentil, Lens culinaris, carbonized seeds from Late Bronze Age Manole, Bulgaria. C–Faba-like bean, Vicia faba var. minor, carbonized seeds from Copper Age Chibanes, Portugal. D–Bitter vetch, Vicia ervilia, carbonized seeds from Late Bronze Age Manole, Bulgaria. E–Chickpea, Cicer arietinum, carbonized seeds from Early Bronze Age Arad, Israel (Zohary and Hopf 1973). Note conspicuous size difference between archaeological and modern seeds; caused by the effect of charring and/or selection under domestication towards larger seeds.

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vetches are climbers with tender branches and characteristic tendrils. Wild lentils are small, delicate plants. Under cultivation, most of these pulses evolved stiffer free-standing, robust stems, and a reduced dependence on climbing—traits that make them better adapted to grow in stands in tilled fields. The wild-type chemical defences have also been selected against. Many wild legumes contain potent toxins and antimetabolites in their seeds to protect them against animal predation. Cultivars frequently lack, or contain only reduced amounts of these toxic compounds. In some others, fermentation (e.g. in soybean) or cooking (e.g. in common bean) is necessary to render the seeds safe for human consumption.

Lentil: Lens culinaris Lentil ranks among the oldest and the most appreciated grain legumes of the Old World. It is grown from the Atlantic coast of Spain and Morocco in the west, to the Indian subcontinent in the east (Smartt 1990; Zohary 1995a). In Mediterranean grain agriculture, it is a characteristic companion of wheat and barley. Compared to the cereals, yields are relatively low, but lentil stands out as one of the most nutritious and tasty pulses. The protein content is about 25%, and lentil constitutes an important meat substitute in peasant communities. Large quantities of lentils are produced and consumed (in soup, paste, in mixture with wheat or rice) in south-west Asia, India, Pakistan, Ethiopia, and, countries bordering the Mediterranean Sea. The domesticated L. culinaris subsp. culinaris Medik. [= Lens culinaris Medik.] manifests a wide range of morphological variation both in its vegetative and reproductive parts. Like many other annual grain crops, lentil is predominantly self-pollinated. Consequently, numerous true breeding lines and aggregates of land races have evolved in this grain crop. All cultivated lentil varieties are diploid (2n = 2x = 14 chromosomes) and predominantly interfertile. Conventionally, lentil cultivars are grouped in two intergrading clusters of seed sizes: (a) smallseeded lentils (subsp. microsperma), with small pods and small 3–6 mm seed; (b) large-seeded lentils

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(subsp. macrosperma), with larger pods and with seed attaining 6–9 mm in diameter. As in other pulses, domestication brought about the retention of the seed in the pod (pod’s indehiscence) and a gradual increase in seed size. Macrosperma forms are to be regarded as more advanced; they start to appear rather late in archaeological sequences— only in the third millennium BP. Carbonised seed is the main element in archaeological assemblages. Occasionally pods, or fragments of pods, occur as well.

Wild ancestry Cultivated lentil belongs to Lens Mill., a small leguminous genus, now known to contain the following taxa (van Oss et al. 1997; Ferguson et al. 1998, 2000; Sonnante et al. 2009): (i) the crop L. culinaris Medik. subsp. Culinaris; (ii) the well-recognized wild progenitor L. culinaris subsp. orientalis (Boiss.) Ponert [= L. orientalis (Boiss.) Shmalh.]; (iii) two more subspecies of L. culinaris – subsp. odemensis (Ladiz.) M.E. Ferguson et al., and subsp. tomentosus (Ladiz.) M.E. Ferguson et al.; and (iv) additional three ordinary wild species: L. nigricans (M. Bieb.) Godr., L. ervoides (Brign.) Grande, and L. lamottei Czefr. All members of Lens, both wild and domesticated, are annual, ephemeral, diploid, predominantly selfpollinated plants. All the wild taxa seem to be reproductively isolated from one another by various combinations of reproductive isolation barriers, such as the genetic system of selfing, the lack of crossing, the invariability of the F1 inter-taxa hybrids, chromosome behaviour in meiosis, and consequently, inter-specific hybrid sterility. The presence of such barriers indicates that we are confronted with fully developed biological species. In contrast, the cross-fertility of culinaris lines with orientalis ones, indicate that both taxa should be lumped together (as subspecies) in the crop’s biological species (Zohary 1995a). Geographically, the genus Lens is a Mediterraneanbasin, south-western, and central Asian element (Map 8) (Zohary and Hopf 1973; van Oss et al. 1997; Ladizinsky 1993, 1999; Ferguson et al. 1998). The wild progenitor of the domesticated plant, subsp. culinaris, shows close morphological, cytogenetic, and molecular affinities, to wild subsp. orientalis. In

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Map 8 Geographical distribution of wild lentil, Lens culinaris subsp. orientalis [= L. orientalis] (based on Zohary and Hopf 1973; Ladizinsky 1993; Gabrielian and Zohary 2004). Wild lentil extends eastwards beyond the boundaries of this map into north Afghanistan and adjacent central Asia (Ladizinsky and Abbo 1993).

fact, subsp. orientalis (Fig. 23) looks like a miniaturized subsp. culinaris, but bears pods that burst open immediately after maturation. Cytogenetic tests have shown (Ladizinsky et al. 1984) that in southwest Asia, subsp. orientalis is chromosomally variable. In contrast, the populations growing in central Asia are chromosomally uniform and exhibit the ‘standard’ chromosomal type only. It also contains five to six additional chromosomal races that differ from the standard race by one or even two chromosomal rearrangements (mostly translocations). Cytogenetic affinities of the crop with other Lens species are less tight. Crosses between L. culinanis (and/or L. orientalis) with L. odemensis Ladiz. result in viable F1 hybrids. They are totally sterile since their parents differ by four chromosomal rearrangements. The four other Lens species, namely L. nigricans (M. Bieb.) Gord., L. ervoides (Brign.) Grande, L. tomentosus Ladiz., and L. lamottei Czefran. turned out to be genetically more remote. Crosses between them and the crop ended in hybrid embryo break-

down (Ladizinsky et al. 1984; van Oss et al. 1997). Finally, cpDNA tests (Mayer and Soltis 1994), as well as several other molecular comparisons carried out on the various Lens species (for enumeration, consult van Oss et al. 1997), indicated close genetic links between domestic L. culinaris and wild-type L. orientalis. They also revealed much wider genetic distances from the other wild taxa. Ladizinsky (1999) examined cpDNA restriction patterns. This, and additional information from crossability and chromosomal architecture, led him to conclude that Lentil originated from the territories around Turkey and north Syria. The same areas were also mentioned in later rDNA studies conducted by Sonnante et al. (2003) and summarized recently by Sonnante et al. (2009). The accumulated botanical and genetic evidence in Lens establishes orientalis as the wild progenitor of the cultivated lentil. It also indicates from which orientalis chromosome type the crop was derived. As argued by Zohary (1999), the rich chromosomal

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Fig. 23 Wild lentil, Lens culinaris subsp. orientalis [= L. orientalis]—a flowering and fruiting plant (Zohary and Hopf 1973).

polymorphism found in the wild progenitor, compared with the chromosomal uniformity in the cultivars, suggests that this pulse crop was taken into cultivation only once or a very few times. Finally, the botanical naming in Lens needs an adjustment. Since it is now clear that wild L. orientalis is the progenitor stock from which the crop has been derived, this wild lentil should be regarded as part of the crop’s biological species. Its appropriate taxonomic ranking is therefore L. culinaris subsp. orientalis. Like other grain crops, wild lentils differ from the domesticated varieties in their seed dispersal biology. As already mentioned, in orientalis plants the pods burst open upon maturation. In contrast, in cultivars the pods do not dehisce immediately and the seeds are retained. This difference is governed by a single mutation (see Table 7), the non-dehiscent condition (the domestic type) being recessive to the dehiscent one (the wild type). Geographically, orientalis is a south-western and Central Asiatic element (Map 8). It is distributed

over Armenia, Turkey, Syria, Lebanon, Israel, Jordan, north Iraq, west and north Iran, and reappears in Turkmenistan, Tajikistan, and Uzbekistan. Subspecies orientalis grows primarily on shallow, stony soils, and gravelly hillsides in open or steppe-like habitats. It also enters disturbed localities such as stony patches or stone heaps bordering orchards and cereal cultivation. In most parts of its distribution, L. orientalis is rather inconspicuous or even rare. It usually forms small, scattered colonies. However, on stony slopes of Mt Hermon, in the Anti-Lebanon, the oak park-forest belt of southern Turkey, and the western escarpments of the Zagros range, L. orientalis is locally common at 1200–1600 m altitude (Ladizinsky and Abbo 1993). Frequently it grows side by side with bitter vetch (Vicia ervilia).

Archaeological evidence Lentils seem to be closely associated with the start of wheats and barley domestication in south-west

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Asia. Probably, this legume was introduced into cultivation in this region together with emmer, einkorn, and barley; that is, it should be regarded as a founder crop of Old World Neolithic agriculture (Zohary and Hopf 1973). Lentils were apparently collected from the wild in south-west Asia before the firm establishment of farming villages. Wild lentils first appeared in Mousterian, ca. 50,000– 60,000 BP, Kebara Cave, Mt. Carmel (Lev et al. 2005), and Upper Palaeolithic, ca. 23,000 cal BP, Ohalo II (Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004, 2008), Israel. Small, carbonized seeds of lentil have been retrieved from several predomestication sites across the Fertile Crescent: Mureybit, ca. 11,800–11,300 cal BP (van Zeist and Casparie 1968; van Zeist and Bakker-Heeres 1986), Tell Abu Hureyra, ca. 13,400–11,350 cal BP (Hillman 1975, 2000a; Hillman et al. 2001), Djade el Mughara, ca. 10,700–10,400 cal BP (Willcox et al. 2008) in Syria, and from Netiv Hagdud, ca. 11,300–11,100 cal BP (Kislev 1997), Israel. In these early settlements we find the collections of wild L. orientalis together with wild wheats and wild barley, and evidently their pre-domestication cultivation (Weiss et al. 2006; Willcox et al. 2008). A few small lentil seeds have also been found in the Palaeo- and Mesolithic (pre-farming) layers of

Fig. 24 Carbonized seeds of domesticated lentil, Lens culinaris, cotyledons only, no seed coat survived charring, PPNB Yiftah’el, Israel (photograph kindly provided by O. Simchoni).

Franchthi Cave (ca. 15,500–8,750 cal BP), Greece (Hansen 1991a, 1992), and from Early and Middle Neolithic layers in Grotta dell’Uzzo (ca. 7,650–6,450 cal BP), Sicily (Costantini 1989). It is very likely that they represent collected local wild L. nigricans. Somewhat later, at the end of the tenth and in the ninth millennia BP, charred seeds of lentil appear in most of the Pre-Pottery Neolithic B (PPNB) early farming villages in south-west Asia (Map 1). The seeds are still similar in size to those of wild forms (2.5–3.00 mm in diameter), and usually do not occur in quantities. Yet they are always associated with domesticated wheat and barley. Among the richest sites are Tell Aswad, ca. 10,200–9,550 cal BP (van Zeist and Bakker-Heeres 1982 (1985), Tell Abu Hureyra, ca. 10,200–8,700 cal BP (Hillman 1975, 2000a, 2000b, 2001; de Moulins 1997, 2000; Hillman et al. 1989), Jericho, ca. 10,250–9,500 cal BP (Hopf 1983), Çayönü, ca. 10,600-9,900 cal BP (van Zeist 1972; van Zeist and de Roller 1991–2; van Zeist et al. 2003), and Ali Kosh, Iran, ca. 9,600–8,800 cal BP (Helbaek 1969). A large hoard of carbonized lentils was recovered from the ca. 10,100–9,700 cal BP Middle Pre-Pottery Neolithic B Yiftah’el, north Israel (Fig. 24). The size of the hoard (7.4 kg, ca. 1,400,000 seeds) and its contamination by the fruits of the weed Galium tricornutum (which is a characteristic weed in lentil cultivation) indicate that there and then, lentil was already cultivated (Garfinkel et al. 1988). Large amounts of lentil seeds were discovered also in somewhat later phases of the Neolithic settlements in the south-west Asia: in ca. 9,450–9,300 cal BP Jarmo, Iraq (Helbaek 1959a, 1960; Braidwood 1960), in ca. 9,250–9,000 cal BP Tell Ramad, Syria (van Zeist and Bakker-Heeres 1985), in ca. 8,200– 7,800 cal BP ceramic Hacilar (Helbaek 1970), in ca. 7,800–7,350 cal BP Girikihaciyan (van Zeist 1979–80), and in ca. 8,350–7,750 cal BP Tepe Sabz, Deh Luran Valley, Iran (Helbaek 1969). The Tepe Sabz lentils had already attained 4.2 mm in diameter. This is an obvious development under domestication. Lentils appear frequently in Cyprus as early as Aceramic Neolithic: in ca. 9,000–7,600 cal BP Dhali Agridhi, Idalion (Stewart 1974), ca. 8,700–6,800 cal BP Cap Andreas-Kastros (van Zeist, 1981), and ca. 8,400–8,200 cal BP Khirokitia (Waines and Stanley Price 1975–1977; Hansen 1991b, 1994).

PULSES

In the eighth millennium BP, frequent seeds and pods of small-seeded lentil, were found also in ca. 7,950–7,150 cal BP Aratashen and Aknashen, Armenia (Hovsepyan and Willcox 2008). In the eighth and seventh millennia BP, lentils seem to be closely associated with the spread of Neolithic agriculture into south-eastern Europe (Map 2—Plate 6). Here, too, they are found together with domesticated emmer, einkorn, and barley. Charred remains of lentil-seed are present in almost all early Neolithic Greek agriculture settlements such as aceramic Ghediki (Renfrew 1979), the pre-ceramic basal levels of Argissa-Magula (Hopf 1962), ca. 8,650—8,200 cal BP Sesklo (Hopf 1962; Kroll 1981a) and Nea Nikomedeia (van Zeist and Bottema 1971). Lentil remains are also frequent in later Neolithic and Bronze Age contexts in the Greek world, such as ca. 8,650–8,400 cal BP, Knossos (Sarpaki 2009) and second half of fifth millennium BP Kastanas (Kroll 1983, 1984). They were also found in large amounts in some of the Early and Middle (ca. 7,500–6,950 cal BP) Neolithic Bulgarian sites such as ca. 8,000–7,550 cal BP Karanovo Mogila (Thanheiser 1997; Marinova 2004, 2006), as well as in ca. 8,200–6,650 cal BP Anza (Starčevo culture) in Macedonia (Renfrew 1976) and in some Impressed Ware sites in southern Italy (Costantini et al. 1997). In Germany, as well as in several adjacent central European countries, lentil seeds were uncovered in several Linearbandkeramik sites (Willerding, 1980; Körber-Grohne 1987). Such sites are ca. 7,500–6,550 cal BP Aldenhovener Platte (Knörzer 1973, 1974, 1997), ca. 7,150–6,800 cal BP Schletz in Austria (Schneider 1994; Kohler-Schneider 2007), ca. 7,400– 6,550 cal BP Bylany, in the Czech Republic (Tempír 1979), ca. 7,550–6,700 cal BP Menneville, France (Bakels 1984, 1991; Ilett et al. 1995) and also in ca. 7,400–7,050 ca BP Coveta de l’Or and Cova de Cendres, Spain (Hopf and Schubart 1965; Lopez 1980; Buxó 1997), frequently together with pea. In Hungary, lentil occurs in Middle Neolithic, ca. 6,350 cal BP Dévaványa-Réhelyi dűlő (Hartyányi et al. 1968), and in Rumania in Vinča culture Liubcova (Cârciumaru 1996). As in the case of the pea (pp. 85–86) and other pulses, lentils in many Bronze Age settlements in Europe seem to be sparser than in Neolithic times.

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Lentils accompany wheats and barley in the spread of south-west Asian agriculture southwards to Egypt and eastwards to the Caspian basin and the Indian subcontinent. The earliest records of Lens from Egypt come from ca. 7,150–5,950 cal BP Neolithic Merimde (Wetterstrom 1993). Lentils have also been found in pre-dynastic, ca. 6,000–5,500 BP Amratian (Naqada I) culture sites (Wetterstrom 1993), and at sites of all epochs ever since. Further east, charred lentil seeds appear in ca. 2,350–2,100 cal BP Shortughai, Afghanistan (Willcox 1991), and in several Harappan culture sites in Pakistan and in north-west India—this from the older phases of this culture onward (Fuller 2002). Some of the lentil remains discovered in Miri Qalat, south-west Pakistan (Tengberg 1999) might be even older. In analyzing lentil remains obtained from the early south-west Asian sites, it is often difficult to decide whether they represent wild material or domesticated forms. The seed of wild Lens is morphologically very similar to that of the domesticated pulse. The only trait indicating domestication in the archaeological record is the increase in seed size. Yet this process was slow and gradual. The first signs of seed size increase appear as late as in the end of the eighth millennium BP; i.e. some two millennia after the definite establishment of wheat and barley cultivation in south-west Asia. We have, however, some circumstantial evidence suggesting that lentil farming is as old as agriculture itself. As already pointed out, the hoard discovered in ca. 10,100– 9,700 cal BP Yiftah’el indicates treatment as a crop. Indicative are also the data on the spread of Neolithic agriculture into Europe (Zohary and Hopf 1973). Lentils are repeatedly encountered in the early European agricultural settlements of the seventh millennium BP, situated far outside the areas in which either L. orientalis or L. nigricans grow wild. This is a strong indication that lentils were already cultivated in these regions. Such early diffusion, together with the main south-west Asian cereals, could only be the outcome of an even earlier domestication, in the area where the wild progenitor occurs. In summary, archaeological remains do not provide us with any direct diagnostic trait for a conclusive determination of the start of lentil domestication. Moreover, it is very doubtful whether comparative

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morphology will provide us with such clues in the future. Yet once Neolithic agriculture is soundly established, cultivation of lentil is part of it. The available archaeological information on early remains of lentil comes from south-west Asia. This is the very territory over which L. orientalis is geographically distributed.

Pea: Pisum sativum The pea ranks among the oldest grain legumes of the Old World. From its early beginnings, this crop has been a close companion of wheat and barley cultivation (Zohary and Hopf 1973). Pea, Pisum sativum L., is well adapted to both warm Mediterraneantype and cool temperate conditions. In peasant communities in south-west Asia, the Mediterranean basin, temperate Europe, Ethiopia, and north-west India, it constitutes an important source of protein for human consumption. The protein content of the seed is about 22%. Today, pea ranks among the world’s most important pulses (Smartt 1990; Davies 1995). It is grown quite extensively in cool regions such as northern Europe and north-western USA. Mature dry seeds were the principal product in classical times. Today, a substantial proportion of the seed and/or pods are also is harvested as immature vegetable crop either for direct consumption or for canning and freezing. Pea is a diploid (2n = 2x = 14 chromosomes) and predominantly self-pollinated crop. As a consequence of the self-pollination system, variation in pea is moulded in numerous true breeding lines. These were used by Gregor Mendel for his classic genetic experiments, some one hundred and sixty years ago. Cultivated pea shows a wide range of morphological variation. Hundreds of land races are recognized and dozens of modern cultivars have been produced by breeders. All are inter-fertile, or at least partly inter-fertile. Cultivars differ from one another in numerous traits such as the height and habit of the plant (from short field types to tall climbers), the colour of the flower (blue, purple, to white), the size and form of the seed, the texture and the colour of the seed coat, and the colour of the cotyledons (yellow or green). Forms with coloured flowers, relatively small seeds, and long vines are frequently called field peas (var. arvense), while cul-

tivars with white flowers, large seeds, and shorter branches are known as garden peas (var. sativum). As in many other grain legumes, the conspicuous features of evolution under domestication are: (i) indehiscence of the pod—the retention of seed on the ripe pod, a trait governed by a single recessive mutation (Table 7, p. 61); (ii) the gradual increase of seed size from 3–4 mm to 6–8 mm in diameter; and (iii) the reduction of the relatively thick texture and rough surface of the seed coat (testa), which evolved to be thin and smooth, resulting in the breakdown of the germination inhibition of wild peas. Seed remains constitute the bulk of pea material recovered from archaeological excavations. Other plant parts, such as pods, were hardly ever retrieved.

Wild ancestry The domesticated pea belongs to a small leguminous genus, Pisum L., restricted, in the wild, to the Mediterranean basin and south-west Asia. All members of this genus are annual, diploid (2n = 2x = 14 chromosomes) predominantly self-pollinated plants. On the combined basis of morphology, ecology, cytogenetics, and molecular analysis, most botanists now recognize two biological species in Pisum:P. sativum and P. fulvum (Davis 1970; Ben Ze’ev and Zohary 1973; Waines 1975; Waines and Stanley Price 1975–1977). P. sativum L. (the crop complex) contains a variable aggregate of cultivated peas, and numerous wild races closely related to the crop. All forms within the complex readily cross with one another and the hybrids are fully or almost fully fertile. All show full chromosome homology except for a single reciprocal translocation found in some wild forms and also in very few cultivated varieties. P. fulvum Sibth. & Sm. is a distinct east-Mediterranean wild species with characteristic yellow-brownish flowers, and chromosomes that are considerably divergent from those present in P. sativum. The two species are reproductively well isolated in nature, and their hybrids are semi-sterile. The wild forms of P. sativum fall into two main morphological types: (i) A tall ‘maquis type’, previously called P. elatius M. Bieb. [= P. sativum L. subsp. elatius (M. Bieb.)

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Representative sites of wild humble peas Representative sites of wild elatius peas

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Map 9 Geographical distribution of the two main wild races of pea, Pisum sativum: (i) ‘steppe-type’ humile forms, and (ii) ‘maquis-type’ elatius forms (based on Zohary and Hopf 1973). In the west-Mediterranean basin, wild elatius peas extend beyond the boundaries of the map and reach as far as Spain.

Aschers. & Graebn.] is omni-Mediterranean in its distribution and it thrives as a sporadic climber in maquis formations in the relatively mesic parts of the Mediterranean basin. Occasionally elatius peas also colonize hedges and terraces bordering agricultural fields. (ii) A shorter, more xeric ‘steppe type’ (Fig. 25), formally named P. humile Boiss. & Noë [= P. syriacum (Berger) Lehm; P. sativum L. var. pumilio Meikle], is geographically restricted to southwest Asia (Map 9). It occurs in the deciduous oak park-forest belt and in open, steppe-like, herbaceous vegetation, characteristic to southwest Asia, i.e. in the same zone that harbours the wild progenitors of wheats, barley, lentil, and flax. From such primary habitats humile peas spill over to secondary habitats and occasionally infest cereal cultivation. In their general habit, some humile forms resemble the cultivated legume closely, and differ from it mainly by their rough seed coat, and by pods

that burst open and disperse the seed soon after maturation. The boundaries between these two principal wild races are occasionally blurred. Intergrading forms are particularly common, especially in Turkey. Spontaneous hybridization between humile or elatius peas and the domestic varieties also occurs sporadically. Chromosomally, the wild peas of the P. sativum complex fall into two chromosomal types (Ben Ze’ev and Zohary 1973). Two humile collections, one from Mt Hermon area and the other from the environs of Ankara, Turkey, were found to have a chromosome complement identical to that prevailing in the cultivated varieties. Hybrids between such tame and wild forms show normal chromosome pairing and are fully fertile. In contrast, four other humile collections from south and central Israel, and elatius material from Israel and Italy, differ from the crop by a single, chromosomal

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translocation. Hybrids in such cases show full chromosome pairing but some reduced fertility due to the translocation heterozygosity. Nasiri et al. (2009) have examined SSR (microsatellite) markers of wild peas and cultivars. They have found that these markers effectively differentiate between cultivars and the wild accessions. In addition, P. sativum subsp. fulvum was found to be the closest relative of the cultivars, but it should be noted that subsp. humile was not tested in this study. In summary, the available evidence from the living plants points to wild humile peas as the progenitor stock for pea domestication. Humile peas show closer morphological similarities than elatius peas to the domesticated aggregate and grow in steppes or steppe-like habitats; i.e. under open conditions that are not very different from those prevailing under domestication. The Turkish and the south Levant forms, having chromosomes identical with

those present in the cultivars within humile peas, should be regarded as the primary ancestral stock. This is also supported by chloroplast DNA comparisons (Palmer et al. 1985). Yet it is very likely that the humile forms with the chromosome translocation, as well as the more mesic wild elatius peas, contributed some genes to the domesticated ensemble through occasional secondary hybridization. Finally, it is important to note the occasional testa remains in archaeological pea finds. These finds allow us to differentiate between domesticated peas, those with smooth seed coat, and the wildtype seeds, those with rough seed coat (Werker et al. 1979; Butler 2009).

Archaeological evidence Remains of peas are present in many of the PrePottery Neolithic B farming villages that developed in the Fertile Crescent in the second half of the tenth

Fig. 25 Wild pea (‘steppe-type’) Pisum sativum subsp. humile [= P. syriacum]—a flowering and fruiting plant (Zohary and Hopf 1973).

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to the ninth millennium BP (Map 1). The first findings of wild-form pea come from Upper Palaeolithic ca. 23,000 cal BP Ohalo II, Israel (Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004, 2008). Some of the earliest finds of cultivated peas come from ca. 10,500–10,200 cal BP Tell Aswad in south Syria (van Zeist and Bakker-Heeres 1985), ca. 10,250–9,550 cal BP Çayönu in south-east Turkey (van Zeist and Bakker-Heeres 1985), ca. 9,900–9,550 cal BP PPNB Jericho (Hopf 1983) and ‘Ain Ghazal, Jordan (Rollefson et al. 1985). Larger quantities of peas were available from somewhat later Neolithic phases in south-west Asia—from the eighth millennium BP. Carbonized seed accompanied the domesticated wheats and barley in ca. 9,350-8,950 cal BP Çatalhöyük (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007), and a large hoard in final Neolithic, ca. 8,550–8,150 cal BP Erbaba, Turkey (van Zeist and Buitenhuis 1983). Few seeds have been found in ca. 9,450–9,300 cal BP Jermo, Iraq (Helbaek 1959b, 1960; Braidwood 1960). In contrast to the wheats and barley, the earliest archaeological remains of pea do not provide us with simple traits for a foolproof recognition of domestication (Zohary and Hopf 1973). In domesticated peas there is a general trend towards an increase in seed size and hilum elongation, but such changes occurred gradually in the course of domestication. In early finds there is considerable overlapping in the dimensions of wild and domesticated forms. Perhaps the most reliable indication of domestication in peas is provided by the surface of the seed coat (testa)—as long as it is preserved. Wild peas are characterized by a rough or granular surface, while domesticated varieties have smooth seed coats. However, seed coats in most finds are missing, which makes it impossible to ascertain whether they are wild or domesticated. The lower levels, ca. 10,250–9,550 cal BP, of Çayönü (van Zeist and Bakker-Heeres 1985) retained some fragments of seed coats showing a rough surface, thus indicating a collection from the wild. Wild-type seed coats occur even much later in Late Neolithic, ca. 8,200– 7,800 cal BP, Hacilar (Helbaek 1970). Significantly, the remains from upper levels of Çayönü include a single seed with a smooth seed-coat; and those from ca. 9,350–8,950 cal BP Çatalhöyük (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007), Late PPNB 8,700–

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7,000 cal BP Can Hasan I (Renfrew 1968), and from ca. 9,450–8,600 cal BP Bouqras (van Zeist and Waterbolk-van Rooijen 1985) show the smooth seed coat characteristic of domesticated varieties. This strongly suggests that domestication of peas in south-west Asia is as old as the domestication of wheats and barley. During the ninth millennium BP, peas spread outside the Fertile Crescent into Cyprus, the Aegean (Fig. 26), and the Balkan spheres. Peas appear in Aceramic Neolithic Cyprus in: ca. 9,000–7,600 cal BP Dhali Agridhi, Idalion (Stewart 1974), ca. 8,700– 6,800 cal BP Cap Andreas-Kastros (van Zeist 1981), and ca. 8,400–8,200 cal BP Khirokitia (Waines and Stanley Price 1975–1977; Hansen 1991b, 1994). At early Neolithic Greece they were found in ca. 8,400– 8,100 cal BP Nea Nikomedeia (van Zeist and Bottema 1971), and at two early Neolithic, Starčevoculture, sites in Macedonia: ca. 8,200–6,650 cal BP Anza, and ca. 8,150–7,000 cal BP Obre (Renfrew 1974, 1976). The Nea Nikomedeia carbonized seeds are well-preserved and reveal the smooth seed coat of domesticated forms. Peas seem to be associated with the spread of Neolithic agriculture into Europe in the eighth millennium BP (Zohary and Hopf 1973; van Zeis 1980).

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Fig. 26 Charred seed of domesticated pea, Pisum sativum, cotyledons only, no seed coat survived charring, Late Neolithic Dimini, Greece (Kroll 1979).

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Here again they are closely associated with wheats and barley production (Map 2—Plate 6). At the first half of the eighth millennium, peas were found mainly in the southern European countries bordering the Mediterranean, while in the second half they reached the middle European countries. Peas are present in Early Neolithic Bulgaria: ca. 8,000–7,700 cal BP Kovacevo (Popova 1992; Marinova 2006), ca. 7,850–7,700 cal BP Kapitan Dimitrievo (Marinova 2006, forthcoming), and ca. 7,850–7,500 cal BP Ašmaska Mogila (Hopf 1973; Renfrew 1979, Table 7), and also in early Neolithic, Starčevo-Criş culture ca. 7,600–7,500 cal BP Sacarovca, Moldavia (Januševič 1984; Kuzminova et al. 1998). In Italy, early Neolithic peas appear first in the south and later in the north, in ca. 8,000–7,600 cal BP Scamuso (Costantini et al. 1997), ca. 7,750–7,150 cal BP La Marmotta (Rottoli 1993, 2002), and finally in ca. 7,550–6,450 cal BP Sammardenchia (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007). Already in the beginning of the first half of the eighth millennium BP, peas appear further east in ca. 8,000–7,150 cal BP Arukhlo 1 and Arukhlo 2, Georgia (Januševič 1984; Lisitsina 1984; Schultze-Motel 1988a), and further west in ca. 7,950–5,450 cal BP Balma Margineda, Spain (Marinval 1995). Later, they reached ca. 7,550– 6,700 cal BP Menneville, France (Bakels 1984, 1991; Ilett et al. 1995). In the second half of the eighth millennium BP, pea is associated with the spread of the Linearbandkeramik (LBK) culture in Europe (Willerding 1980) as well as the expansion of agriculture to Egypt. In the lower Rhine Valley (Rosdorf), peas are common in LBK sites dated ca. 7,450-7,150 cal BP (Kirleis and Willerding 2008); and in central Germany large amounts of charred seeds have been recovered from similar LBK settlements, such as ca. 6900–6500 cal BP Dresden-Nickern (Baumann and Schultze-Motel 1968). In these samples, seed coats are frequently well-preserved and, with one exception, they show the smooth surfaces characteristic of the domestic crop. Also in LBK Třtice in Bohemia (Tempír 1973, 1979), peas appear in quantities; and they reappear in several early Neolithic sites in Moldavia, like ca. 7,600–7,400 cal BP Sacarovca (Januševič 1984; Kuzminova et al. 1998; Monah 2007b). All over this area, from Ukraine (e.g. Rivne,

Pashkevich 2003) to western Germany, peas become more frequent in late Neolithic times. The earliest finds from Poland, so far, come from Funnel Beaker contexts in ca. 5,600–5,300 cal BP Čmielów (Klichowska 1976). A hoard of pea seeds was found in a Dudéşti culture context in ca. 6,500–6,000 cal BP Cîrcea, Rumania (Cârciumaru 1996). In the west Mediterranean basin, a few seeds have been found in ca. 7,950–5,450 cal BP Balma Margineda, Spain (Marinval 1995), and the evidence continues to the Chalcolithic and Bronze Age layers. Generally, finds of peas, as well as of other pulses, are sparser throughout European Bronze Age compared to the Neolithic finds. Richer deposits of pulses, including lentil and broad bean, occur again in the European settlements of later periods (Zohary and Hopf 1973). Such finds came from ca. 4,150–3,200 cal BP Canton of Grisons, Switzerland (Jacomet et al. 1998, 1999), ca. 4,950–3,350 cal BP Albertfalva, Hungary (Gyulai 2003), ca. 3,200–2,700 cal BP Stillfried, Austria (Kohler-Schneider 2003, 2007), ca. 3,360-3,700 cal BP Wange and Overhespen, Belgium (Bakels 1992), and ca. 3,900–3,350 cal BP Nitriansky Hrádok, Czech Republic (Tempír 1969; Kühn 1981). Pea remains appear also in late eighth millennium BP Neolithic settlements of the Nile Valley, Caucasia and Transcaucasia. In Egypt, early finds come from ca. 7,150–5,950 cal BP Merimde, Nile delta (Wetterstrom 1993) and from ca. 5,650–5,300 cal BP pre-dynastic Nagada settlements, Upper Egypt (Wetterstrom 1993). Further east, the earliest finds are of a later date. Pea remains were retrieved from ca. 2,350–2,100 cal BP Shortughai, Afghanistan (Willcox 1991). They are present in ca. fifth millennium BP Harappa (Vishnu-Mittre and Savithri 1982), and in several other Harappan sites in Pakistan and north-west India, from the older phases of this culture onward (Fuller 2000). In conclusion, the archaeological evidence establishes pea as one of the founder grain crops of south-west Asian Neolithic agriculture. Since this early start, pea seems to be a consistent element in Neolithic and Bronze Age food production throughout West Asia and Europe and a common companion of wheats and barley. The evidence from the living plants complements the

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archaeological finds. The wild humile forms, with chromosomes and cpDNA identical to those prevailing in the cultivated crop, should be regarded as the closest wild stock from which this pulse crop evolved.

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the gradual increase in seed size from 3.5 to 6.0 mm and more. Large-seeded forms are obviously more advanced. Another change is the development of a smooth seed coat, and the reduction of its thickness. Seed remains are the only material recovered todate in archaeological excavations.

Chickpea: Cicer arietinum Chickpea is a valued seed legume of traditional agriculture in the Mediterranean basin, western Asia, as well as India and Ethiopia. It is a member of the grain ensemble found in south-west Asian Neolithic and Bronze Age remains. Chickpea is adapted to a subtropical or Mediterranean-type climate; it grows almost exclusively in the post-rainy season on moisture stored in the soil. The main growers today are in the Indian subcontinent, the countries around the Mediterranean basin and Ethiopia (http://faostat.fao.org). Like lentil and pea, chickpea, with a seed protein content of some 2%, constitutes an important meat substitute in peasant communities. Domesticated chickpea, Cicer arietinum L. subsp. arietinum, is a predominantly self-pollinated annual crop with pods containing one to two seeds. The cultivars show a wide range of variation in size, colour, and shape of the seed, and in the size and form of leaves and flowers. All cultivated varieties are diploid (2n = 2x = 16 chromosomes) and interfertile. Chickpea land races are grouped into two inter-connected clusters (Smithson et al. 1985; Smartt 1990). Large-seeded varieties (known as ‘Kabuli’ type) with relatively smooth, rounded, light-coloured seed coats and pale cream flowers, predominate in the Mediterranean countries and south-west Asia. Varieties producing small, wrinkled seed (‘Desi’ type) with dark-coloured seedcoats and usually purple flowers prevail in the eastern and southern parts of the distribution area of the crop; i.e. in India, Afghanistan, Pakistan, and Ethiopia. Seed dispersal in wild chickpea is attained in two ways: (i) shattering of the seeds from the dry, splitting pods, still attached to the mother plant; and (ii) dispersal of whole inflated pods by wind. As in most other seed legumes, the conspicuous features of evolution under domestication in chickpea are the retention of pods and seeds on the plants and

Wild ancestry The domesticated chickpea, Cicer arietinum subsp. arietinum, is a member of a leguminous genus comprising some forty species, centred in central and western Asia (van der Maesen 1972, 1987; Cole et al. 1998). The majority of Cicer species are herbaceous perennials or dwarf shrubs; some are annuals. The domesticated pulse shows close morphological affinities (and an almost identical seed-protein profile) to two wild species of chickpea: C. echinospermum Davis and C. reticulatum Ladiz. (Ladizinsky and Adler 1976). These two wild chickpeas are diploid, self-pollinated annuals, recorded only in south-eastern Turkey. Externally, they resemble each other closely, but they can be distinguished from one another by testa (seed-coat) texture, which is echinate in C. echinospermum and reticulate in C. reticulatum. Also, they differ from one another by their soil preferences; the first grows mainly on basaltic soils, while the second thrives on limestone bedrock. Of the two wild chickpeas only C. reticulatum (Plate 10) is fully inter-fertile and contains chromosomes homologous to those found in the crop (Ladizinsky and Adler 1976). Crosses between the domesticated chickpea and C. echinospermum, and also those between C. reticulatum and C. echinospermum are more difficult to achieve, and the interspecific F1 hybrids obtained proved to be highly sterile. Attempts to cross the domesticated chickpea with other closely related wild Cicer species— C. pinnatifidum Jaub. & Spach, C. judaicum Boiss., and C. bijugum Rech.—failed altogether, indicating that these wild species are not closely related to the crop. The distribution of C. reticulatum is currently restricted to south-eastern Turkey (Map 10). LevYadunet al. (2000) argue for this limited area, where other wild progenitors grow as well, to be the ‘core area’ for all founder crops. Since this distribution is so limited, molecular research has been conducted on material collected

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100 200

200 miles 400 km

Map 10 Geographical distribution of wild chickpea, Cicer arietinum subsp. reticulatum [= C. reticulatum] (based on Ladizinsky and Adler 1976; Tanno and Willcox 2006b; and unpublished data of D. Zohary).

from this area only. Indeed, the molecular evidence seems to supports the identification of C. reticulatum as the wild progenitor. Both Nguyen et al. (2004), upon examining 214 AFLP loci of 94 different accessions of chickpea and its wild relatives, and Sethy et al. (2006) upon examining microsatellite markers, pointed out the same wild progenitor. In summary, the morphological, cytogenetic, and molecular evidence currently available, points to C. reticulatum as the wild progenitor of the domesticated plant. Consequently, it is referred to as C. arietinum subsp. reticulatum. The central part of the Fertile Crescent seems to be the territory in which chickpea could have been taken into domestication. It is noteworthy that unlike wild C. reticulatum, which is an annual plant that germinates in the rainy season, south-west Asian domesticated

chickpeas are grown as summer crops. This shift to summer growing, is attributed by Abbo et al. (2003) to the interaction with the pathogen Didymella rabiei, the casual agent of Ascochyta blight. Another attraction for the grower to turn chickpea into an early summer crop are the many hours of daylight during the long, summer days, which enhance speedy maturation of seeds, even in colder environments (ibid). It seems that the available potential of chickpea as a major grain crop is not yet fully utilized.

Archaeological evidence Like lentil and pea, chickpea seems to have been closely associated with the start of agriculture in south-west Asia. However, in Pre-Pottery Neolithic

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B contexts in this core area, chickpea is much rarer than the two other pulses. The available evidence indicates that several closely related annual wild Cicer species co-exist in the south-west Asia. A few charred chickpea seeds were recovered from various phases of aceramic Çayönü, ca. 10,250–9,550 cal BP and later (van Zeist 1972; van Zeist and de Roller 1991–2, 2003), from contemporary Nevali Çori (Pasternak 1998), and from ca. 10,200–9,500 cal BP Aşikli Höyük (van Zeist and de Roller 1995) in Turkey. A few more were found in Middle PPNB, ca. 9,400–7,000 cal BP Tell Abu Hureyra, northern Syria (Hillman 1975, 1989; de Moulins 2000). The seeds still correspond in size to those of C. reticulatum and, because these sites are situated within or close to the present distribution area of the wild progenitor, the finds could represent either wild or domesticated material. Chickpea seeds were retrieved from PPNA, ca. 10,600–10,250 cal BP, Tell el-Kerkh, north-west Syria (Tanno and Willcox 2006b). A few of these seeds are somehow larger and plumper than the wild progenitor. Tanno and Willcox (2006b, p. 200) argue that this might indicate that they are ‘intermediate stage’ between wild and domesticated chickpeas. The seeds retrieved from Pre-Pottery Neolithic B, ca. 9,900–9,550 cal BP, Jericho (Hopf 1983) and ‘Ain Ghazal (Rollefson et al. 1985), and those from the Late PPNB level in Ramad near Damascus (van Zeist and Bakker-Heeres 1985) probably represent domesticated forms. The latter sites lie far away from the territory of the wild progenitor, and the specimens from Jericho seem to have smooth coats. Likewise, some 500 chickpeas seeds, few of them with smooth testa, from ca. 8,350–8,050 cal BP pottery Neolithic site of Höyücek, south-west Turkey, positively represent domesticated chickpea (Martinoli and Nesbitt 2003). Early Bronze Age remains of chickpeas are more abundant. Considerable amounts of charred, wellpreserved, fairly large seeds with smooth coats were retrieved in Israel and Jordan from Early Bronze Age Arad (Hopf 1978c), contemporary Jericho (Hopf, 1983), Bab edh-Dhra (McCreery 1979) and ca. 4,900 cal BP Tell es-Sa’idiyeh (Cartwright 2003; and persoal communication). These finds provide a clear indication of chickpea domestication since the only chickpea growing

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wild in Israel and Jordan, namely Cicer judaicum, has significantly smaller seed and a characteristic rough seed coat. A single, well-preserved chickpea was retrieved from a pre-Sesklo layer, eighth millennium BP Otzaki, Greece (Kroll 1981a). A few seeds were also found in Early Neolithic, ca. 8,000–7,700 cal BP Kovacevo (Popova 1992; Marinova 2006) and ca. 7,850–7,700 cal BP Kapitan Dimitrievo (Marinova, forthcoming) in Bulgaria. Richer finds are available from late Neolithic, ca. 6,650–6,300 cal BP, Dimini, Greece (Kroll 1979). They measure 4.24 × 3.77 × 3.51 mm. The domesticated pulse must therefore have reached Europe with one of the first waves of migration of early grain crops from south-west Asia. The crop is missing in Neolithic layers of former Yugoslavia, and the western, central, and northern European countries. From Bronze Age to classical times, chickpea evidently ranked among the popular pulses of the Mediterranean basin and the southwest Asian countries. The only early chickpea find from Egypt is ca. 1,325 BC Tutankhamun tomb (Germer 1989a; Hepper 1990; de Vartavan et al. 2010). In conclusion, the evidence from the living plants and the plant remains discovered in archaeological excavations indicate that Cicer arietinum belongs to the Early Neolithic grain crop assemblage of southwest Asia. Archaeological data is limited, but in this legume, the delimitation of the place of origin is relatively simple: the wild progenitor of the domesticated chickpea is endemic to the central part of the Fertile Crescent. Very probably, this pulse was first brought into domestication there.

Faba bean: Vicia faba Together with lentil, pea, and chickpea, the faba bean, Vicia faba L. (broad bean, horse bean), belongs to the principal pulses of Old World agriculture. It grows well in both warm, summer-dry Mediterranean-type environments, and in the more northerly temperate parts of Europe and Asia (Bond et al. 1985). The erect, robust plant readily bears threshable pods and relatively large seed (Plate 11) with high (about 2–25%) protein content. In some Asian and Mediterranean countries, particularly in Egypt, the dry seeds of the faba bean provide a staple

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protein source for the poor. Unripe pods are also used when green. In both Europe and Asia, the seed is regarded as a valuable animal feed, hence the name ‘horse bean’. Large quantities of V. faba are produced in several Mediterranean countries: Egypt, Morocco, Spain, Italy, Turkey, as well as in Ethiopia, temperate Europe, south-western Asia, and especially in China. Vicia faba is an annual, diploid grain crop (2n = 2x = 12 chromosomes). In contrast to the majority of the grain crops, the faba bean is not fully a selfpollinated crop (Lawes 1980). It is self-compatible, and a considerable amount of cross-pollination takes place in numerous cultivars. Furthermore, when subjected to selfing, many domesticated varieties manifest various degrees of inbreeding depression. On the other hand, some numerous faba cultigens, particularly paucijuga forms from India and Afghanistan, are able to self-pollinate. Under domestication, faba beans developed a considerable amount of morphological variation and different ecological adaptations. Conspicuous intra-specific differences are displayed in various traits like vegetative habit, pod structure, pod shattering, and the shape, colour, and size of the seed. Seed size has served as the principal character for intra-specific subdivision. Faba bean taxonomists (Muratova 1931; Hanelt 1972) recognize three or four intergrading and inter-fertile main types of cultivar forms in V. faba. Forms with relatively small rounded seed measuring 6–13 mm are placed in V. faba var. minor. Some small-seeded forms, grown mainly in India, Pakistan, and Afghanistan, also show reduced numbers of leaflets (mostly one pair per leaf), and have very small flowers. They are frequently regarded as a distinct intra-specific taxon: var. paucijuga. Forms with medium-sized seed, that are 15–2 mm long, 12–15 mm wide, and 5–8 mm thick, are placed in var. major. Large-seeded faba seem to have evolved under domestication relatively late. Archaeological remains ranging from the Neolithic period to Roman times are almost all within the range of var. minor.

Wild ancestry The wild ancestor of the domestic faba bean has not yet been discovered despite intensive research in

the last fifty years. In its general morphological appearance, this domestic pulse resembles another group of large-seeded south-west Asian and Mediterranean wild vetches, namely: V. narbonensis L., V. serratifolia Jacq., V. johannis Tamamsch., V. galilea Plitm. & Zoh., V. hyaenis-ciamus Mouterde, and V. kalakhensis Khattab, Maxted & Bisby. The first three taxa are widely distributed over south-west Asia and the Mediterranean basin, V. galilea is a south-west Asian element, and the last two taxa are strict endemics confined each to a small territory in Syria (Maxted 1995). The morphological similarities between the domestic faba bean and this group of wild vetches are considerable. Consequently, taxonomists dealing with the genus Vicia, used to place both the crop and these wild taxa in a common section (Sect. Faba Aschers. & Graebn.). The ancestry of the crop was sought among these wild taxa. However, extensive cytogenetic tests proved that the faba crop is unique both chromosomally and reproductively. The first surprise was that while V. faba has 2n = 2x = 12 chromosomes, all the abovementioned wild vetches were found to have 2n = 2x = 14 chromosomes. Furthermore, largescale attempts to cross the wild vetches with the faba crop (Schäfer 1973; Ladizinsky 1975) failed altogether, indicating that V. faba is reproductively strongly isolated from these vetches. These finds ruled out the candidacy of the above-mentioned faba-resembling wild vetches for the ancestry of the crop. They are now regarded as a separate aggregate of species ‘Vicia narbonensis complex’, (see Schäfer 1973; Bennett and Maxted 1997). Maxted (1995) went a step further. He separated this 2n = 2x = 14 aggregate from V. faba and considers it as an independent taxonomic section (Sect. Narbonensis (Radzhi) Maxted). Finally, molecular comparisons (van de Ven et al. 1993; Jaaska 1997a; Potokina et al. 1999) also indicate that V. faba is not close to the V. narbonensis complex. In summary, in the case of the faba bean, we still need to discover an elusive 12-chromosomed wild progenitor that, in all likelihood, is very limited in its geographic distribution. There is also the possibility that this wild ancestor is extinct. While these faba-resembling vetches have been ruled out as candidates for the wild ancestry of the crop, their common occurrence in south-west Asia

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and the Mediterranean basin affects our ability to identify V. faba remains in archaeological excavations. In many places today, these wild vetches are locally common. They constitute an attractive element for collection from the wild, and as Enneking and Maxted (1995) argue, V. narbonensis even show some signs of independent domestication. Therefore, one can expect that seed remains of members of the narbonensis group could occasionally appear in archaeological contexts, particularly in the early sites. Distinguishing between the faba crop and these vetches is another story. Considerable similarity exists—both in shape and size—between the seeds of V. narbonensis (or some of its close relatives), and those produced by small-seeded faba cultivars. For these reasons, the separation of V. faba from V. narbonensis complex, in archaeological remains, might be difficult; more so, since charring usually destroys the seed coats. Identification becomes very unreliable when only few seeds (or seed fragments) are available. Unfortunately such paucity characterizes almost all finds retrieved from early archaeological contexts. As a result, archaeobotanical faba remains should not be regarded as ‘V. faba’ but have to be considered as V. faba-like’.

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Israel. Here, hoards of some 2,600 charred seeds (without seed coats) were retrieved (Kislev 1985). The seeds are small (5.5 × 4.7 × 4.0 mm), conforming in size with those of wild V. narbonensis or V. galilea—which are common plants in Yiftah’el area. But compared to the more globular shape of the seeds of these wild vetches, they are flatter and somewhat thicker at the hilum’s side. The last two features suggest that Yiftah’el’s remains belong to V. faba. Yet, this evidence is not conclusive. Apart from this single find, all other early records of V. faba-like remains are indefinite. A few faba-like seeds are available from several Pre-Pottery Neolithic B farming sites in south-west Asia, such as ca. 9,900–9,550 cal BP Jericho (Hopf 1983), Tell Abu Hureyra (Hillman 1975; de Moulins 2000), Cap Andreas-Kastros (van Zeist 1981), or Nevali Çori (Pasternak 1998). Already during the Neolithic, faba-like beans appear also in several southern European countries: in ca. 7,500–6,550 cal BP Scamuso, Italy (Costantini et al. 1997), and in south-eastern Spain, like ca. 7,400–7,050 cal BP Coveta de l’Or (Hopf and Schubart 1965; Lopez 1980), and adjacent Cova de

Archaeological evidence As already noted, the identification of V. faba-like remains from early sites is complicated. All wild progenitors closely resemble (morphologically, as well as at the chromosome and molecular levels) their domestic forbears, but the situation is even more problematic in V. faba, as we do not know its wild progenitor. Earliest charred V. faba-like seeds appear in ca. 13,150–11,250 cal BP Iraq ed Dubb, Jordan (Colledge 2001). However, it is more likely these are either wild or intrusions. Later, during the PPNA, faba-like seeds were found in ca. 10,700– 10,400 cal BP, Djade el Mughara (Willcox et al. 2008), and the rich (437 seeds) find in ca. 10,600–10,250 cal BP, Tell el-Kerkh, north-west Syria (Tanno and Willcox 2006b). A few of these seeds are somehow flat and wedge-shaped, and probably closely resemble the wild progenitor (Tanno and Willcox 2006b, p. 203). The only rich find available so far comes from Middle PPNB, ca. 10,100–9,700 cal BP, Yiftah’el,

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Fig. 27 Carbonized seeds of faba-like bean, Vicia faba var. minor, cotyledons only, no seed coat survived charring, Early Bronze Age Zambujal, Portugal (Hopf 1981).

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Cendres (Buxó 1997). During the seventh millennium it appears in ca. 6,500–5,500 cal BP Kaf That el-Ghar, Morocco (Ballouche and Marinval 2003). But these few, testa-less remains do not permit us to decide whether they represent faba bean or members of the V. narbonensis group. Similarly, the rare, small, globular V. faba-type seeds reported from Greek Late Neolithic, ca. 6,650–6,300 cal BP, Dimini (Kroll 1979) and Sesklo (Hopf 1962, 1981a), also conform well with V. narbonensis group. V. faba appears in several European Bronze Age sites (Fig. 27): ca. 4,950–3,350 cal BP Albertfalva, Hungary (Gyulai 2003), ca. 4,150–3,200 cal BP Canton of Grisons (Graubünden), Switzerland (Jacomet et al. 1998, 1999), ca. 3,200–2,700 cal BP Late Bronze Age Stillfried, Austria (Kohler-Schneider 2001, 2003), and Late Bronze Age, ca. 2,905–2,869 cal BP, Grésine, France (Bouby and Billaud 2001). Significantly, many of the sites north of the Alps are outside (or almost outside) the distribution range of the members of the V. narbonensis group, and this fact minimizes the possibility that the excavated seeds belong to them. Furthermore, seed size in several locations is already larger than that found in the wild vetches. This strongly suggests that we are confronted with domesticated V. faba. Numerous finds of faba beans are available from Iron Age and classical times in Europe and west Asia, indicating that V. faba had been used as a major food source. When the information from the living plants and from the archaeological remains is put together we are led to the following conclusions: (i) The wild progenitor of the domesticated faba bean has not been satisfactorily identified. We still need to discover the elusive 12-chromosome ancestor. (ii) We still know very little about the beginnings of V. faba domestication. A main reason being that the early (PPNA/Early PPNB—twelfth– eleventh millennia BP) finds identified as V. faba seed might actually belong either to the crop or to members of the V. narbonensis species complex. More definite conclusions can only be reached after additional well-preserved archaeological evidence is recovered or when the wild progenitor of the crop is found.

Bitter vetch: Vicia ervilia Bitter vetch, Vicia ervilia (L.) Willd., is a small, annual diploid (2n = 2x = 14 chromosomes), self-pollinated pulse with beaded pods and characteristic angular seeds. At present it is grown as a minor crop in the Mediterranean basin and south-west Asia. The principal producer is Turkey. As its name implies, its seeds are bitter and toxic to humans. However, soaking, leaching, and steaming in water can remove the poisonous substances, making them palatable (van Zeist 1988; Enneking 1995; Miller 2002). Roman literature reports that in Classical times, this vetch was utilized primarily as an animal feed since it is regarded as inferior for human consumption and is only eaten by the very poor, or in times of famine. Pliny the Elder (Natural History: 18.38) states that bitter vetch (‘ervi’) has medicinal properties, like vetch (‘vicia’). He mentions that bitter vetch ‘cured’ the Emperor Augustus. The wild ancestry of the domesticated bitter vetch is satisfactorily established (Zohary and Hopf 1973; Ladizinsky and van Oss 1984). Truly wild forms of this vetch, growing in primary habitats, are restricted in their distribution and known from Anatolia, Armenia, north Iraq, the Anti-Lebanon (including Mt Hermon), and from Jebel Druz (Map 11). They show a striking morphological resemblance to the domesticated crop but differ from it by their dehiscent pods and slightly smaller seeds. Hybrids between the cultivars and the wild forms are fully fertile. Weedy races and feral forms of V. ervilia occasionally infest grain crops and edges of cultivated fields throughout south-west Asia and Greece. Unfortunately, we did not find molecular research aiming to support the identification of the wild progenitor and whether it had a monophyletic or polyphyletic origin.

Archaeological evidence The earliest finds of bitter vetch appear in Mousterian, ca. 65,000–48,000 cal BP, Kebara Cave, Israel (Lev et al. 2005). Afterwards, it was found in several Syrian sites. First in Epi-Paleolithic, ca. 13,400–11,350 cal BP, Tell Abu Hureyra (Hillman 1975; Hillman et al., 1989, 2001), and ca. 11,800–

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200 miles 400 km

Map 11 Geographical distribution of wild bitter vetch, Vicia ervilia (based on Zohary and Hopf 1973; Townsend 1974; Ladizinsky and van Oss 1984; Gabrielian and Zohary 2004).

11,300 cal BP Mureybit (van Zeist and Casparie 1968; van Zeist and Bakker-Heeres 1986), and later in PPNA, ca. 11,500–11,000 cal BP, Jerf el-ahmar (Willcox 2002; Willcox et al. 2008, 2009), and ca. 10,700–10,400 cal BP Djade el Mughara (Willcox et al. 2008). Later, seeds of bitter vetch first appear in several agricultural settlements in Turkey. Thousands of carbonized seed of V. ervilia were found in various phases of Aceramic, ca. 10,250– 9,550 cal BP, Çayönü (van Zeist 1972; van Zeist and de Roller 1991–2, 2003), and in ca. 10,100–9,450 cal BP Aşikli Höyük (van Zeist and de Roller,1995). Bitter vetch is also common in aceramic, ca. 9,450– 8,450 cal BP, Can Hasan III (Hillmam 1972, 1978). In addition, it appears in contexts of ca. 9,350–8,950 cal BP Çatalhöyük (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007), ca. 8,550–8,150 cal BP Erbaba (van Zeist and Buitenhuis 1983), and ca. 8,200–7,800 cal BP Hacilar (Helbaek 1970). In addition V. ervilla was very important in the early phase, ca. 8,500–8,000 cal BP, of Gritille (Miller 2002), and ca. 7,800–7,350 cal BP Girikihaciyan (van Zeist 1979–80). A few seeds, and even a single pod remain, were found in

ca. 7,950–7,150 cal BP Aratashen and Aknashen, Armenia (Hovsepyan and Willcox 2008). However, it is impossible to determine definitely whether these remains represent wild or domesticated material. Later finds, such as Middle Bronze Age Shiloh (Kislev 1993) and Beit She’an (Simchoni et al. 2007), Israel, are also relatively few. Considerable amounts of charred seed of V. ervilia have been discovered in late Neolithic and Bronze Age Greece (Fig. 28). The earliest rich (we refer to the large quantity as indication for the domestication status of the find) and well-identified bitter vetch finds come from ca. 8,400–8,100 cal BP Nea Nikomedeia (van Zeist and Bottema 1971), followed by several late Neolithic and early Bronze Age finds (Renfrew 1979; Kroll 1983). Bulgarian sites show similar situation. Large amounts of bitter vetch were found in early Neolithic, ca. 8,000–7,550 cal BP Karanovo, followed by storages of it in Middle Neolithic, late Neolithic, Late Eneolithic, and Early Bronze Age strata (Thanheiser 1997; Marinova 2004, 2006). Huge, pure hoards of bitter vetch grains were discovered in Middle Neolithic, ca. 7,450–7,300 cal

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5 mm

0

Fig. 28 Carbonized seeds of bitter vetch, Vicia ervilia, cotyledons only, no seed coat survived charring, Late Neolithic Dimini, Greece (Kroll 1979).

BP, Azmaška Mogila (Hopf 1973a; Renfrew 1979). Carbonized remains of this pulse are also very frequent in Bulgaria throughout the Eneolithic and the Bronze Age (Januševič 1978; Renfrew 1979) and sometimes constitute the main plant material retrieved from the sites. Outside Turkey, Greece, and Bulgaria, remnants of the crop are much less common. Some seeds were found in ca. 7,550–6,450 cal BP Middle Neolithic Sammardenchia, Italy (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007), and contemporaneous Grotta dell’Uzzo, Sicily (Costantini 1989). However, over large areas in West Europe and the Mediterranean Basin they are missing altogether. A few grains of bitter vetch were retrieved from Late Neolithic Gomolava, in Serbia (van Zeist 1975). Several finds are reported from Rumania, like ca. 6,350–6,150 cal BP Tell Hârşova (Cârciumaru 1996; Monah 2002; Monah and Monah 2008), including an almost pure hoard from Gumelnitsa culture in Cǎscioarele (Cârciumaru 1996). A few seeds were uncovered in Tripolye culture sites in Moldavia and the Ukraine (Wasylikowa et al. 1991). Some rare finds were also discovered in ca. 5,650–5,300 cal BP

Naqada, Egypt (Wetterstrom 1993). Later, small amounts of bitter vetch have been found in ca. 3,350–3,250 cal BP Bölcske-Vörösgyír, Hungary (Berzsényi and Gyulai 1998), ca. 3,200–2,700 cal BP Stillfried, Austria (Kohler-Schneider 2001, 2003), and ca. 2,905–2,869 cal BP Grésine, France (Bouby and Billaud 2001). Yet on the whole, remains of V. ervilia in Neolithic and Bronze Age contexts show a distinct regional pattern. The cultivation of this pulse in the past seems to have been heavily centered in Anatolia and the Balkans. The combined evidence from the living plants and from the archaeological remains indicates that bitter vetch was a founder crop, part of south-west Asian Early Neolithic crop assemblage. This pulse was probably taken into domestication in Anatolia or the Levant; i.e. in the general area in which it still grows wild today. However, since there are no reliable diagnostic traits by which wild and domesticated forms of bitter vetch (in archaeological remains) can be distinguished from each other, the early archaeological finds could be either. The size and purity of the numerous samples of this pulse retrieved from the Neolithic and Bronze Age sites in

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the Balkans and Turkey, strongly suggests that V. ervilia was domesticated at that time and place. We know very little about the mode of utilization of this bitter seeded legume by the Neolithic and Bronze Age farmers.

Common vetch: Vicia sativa Common vetch, Vicia sativa L., is another member of the genus Vicia that is frequent in Mediterranean grain agriculture. This vetch is very variable, including wild forms, weeds, and cultivars. In modern times, it constitutes a minor crop cultivated for hay and for seeds, grown almost exclusively for animal feed. Similar to bitter vetch, the seeds are not attractive for human consumption. Common vetch is also a frequent contaminant of lentil and bitter vetch cultivation. Seeds of the latter pulses sold in local markets frequently contain scattered V. sativa seed. The domesticated V. sativa is a self-pollinated, diploid plant (2n = 2x =12 chromosomes) with straggling or ascending growth habit and rounded, somewhat compressed, smooth seeds (4.5–7.0 mm in diameter). The cultivars are closely related to an extraordinarily variable (and chromosomally complex) aggregate of wild types and weedy forms, the distribution of which is centred in the Mediterranean basin (Zohary and Plitman 1979; Maxted 1995). All are now grouped, together with the cultivars, in the V. sativa complex. Most cultivars, together with morphologically closely related weeds and escapees are placed in V. sativa subsp. sativa.

Archaeological evidence Carbonized seeds of V. sativa have been reported from several Neolithic and Bronze Age sites in south-west Asia and Europe. Since seed sizes of weedy forms and wild types overlap those found in the cultivars, it is difficult to conclude whether the remains represent domesticated forms, weedy contaminants or collection from the wild. Definite indications of sativa cultivars are available only from Roman times onward. The earliest archaeological records of V. sativa comes from the Natufian and Neolithic Tell Abu Hureyra, Syria (Hillman 1975, 2001; Hillman et al. 1989; de Moulins 2000) and from pre-ceramic

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Neolithic Can Hasan III, Turkey (Hillman 1972). They are followed by several records from Neolithic and Eneolithic Bulgaria (Renfrew 1973), Italy (Rottoli 1993; Pessina and Rottoli, 1996; Rottoli 2002, 2005; Rottoli and Pessina 2007), Hungary (Hartyáni and Nováki 1975), and Slovakia (Hajnalová, 1989). Common vetch is also reported from Predynastic Maadi, Egypt (van Zeist and de Roller 1993, 2003), and from several Bronze Age contexts such as the second half of the fifth millennium BP Ak-Tepe near Ashkabad, Turkmenia (Lisitsina and Prishchepenko 1977, as cited by Schultze-Motel 1974b), and from Slovakia (Kühn 1981).

Grass pea: Lathyrus sativus Grass pea or chickling vetch, Lathyrus sativus L., is a minor pulse crop of traditional agriculture in the Mediterranean basin, south-west Asia, Ethiopia, and the north-western parts of the Indian subcontinent (Kearney and Smartt, 1995). India is the main producer today, and there L. sativus is appreciated for its ability to grow in dry places and on poor soils. Grass pea is a diploid (2n = 2x = 14 chromosomes), predominantly self-pollinated annual with branched, straggling, or climbing habit, blue (sometimes violet or white) flowers and characteristic smooth seeds with pressed sides. Recent cytogenetic studies found out a mixed mating system with an outcrossing rate of 36% (Gutiérrez-Marcos et al. 2006). At present the pulse is used mainly as animal feed, though in India and Ethiopia it serves also as an article of human diet and is consumed by the very poor, and in times of famine. For this reason, as well as its palatability, L. sativus is currently examined for serving as a crop replacement, especially in depleted fields (Polignano et al. 2009). Consuming large amounts of grass pea seed can be dangerous. The seed contains a water-soluble non-protein neurotoxin amino acid that causes Lathyrism, a crippling neurological disorder. Because the neurotoxin is a water-soluble, one hour boiling of the seeds removes some 70% of it (Jha 1987). Domesticated grass pea shows close morphological resemblance to a group of wild Lathyrus species distributed over the Mediterranean basin and southwest Asia. It seems closest to L. cicera L., a wild grass

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pea growing in several eastern Mediterranean and south-west Asian countries (Greece, Turkey, North Iraq, North Iran, Transcaucasia, and Egypt). In these countries, L. cicera abounds as a weed in cereal cultivation. L. cicera can be crossed with L. sativus, and the hybrids produced are at least partly fertile (Kearney and Smartt 1995). Therefore, L. cicera could be considered as the wild progenitor from which the domesticated grass pea was derived. But L. cicera is very variable, and is cytogenetically complex. Moreover, several other Lathyrus species also show close morphological affinities with L. sativus. Therefore, the final determination of ancestry in this crop has to depend on critical cytogenetic and molecular tests. These are as yet unavailable. As in many other grain legumes, domestication of the grass pea resulted in retention of the seeds in the pods, loss of seed dormancy, and an increase in seed size. Seeds of L. sativus cultivars are somewhat larger (6–8 mm in diameter) than those of their wild relatives (5–6 mm). Unfortunately, we did not find any molecular research seeking to support the identification of the wild progenitor and whether it was a monophyletic or polyphyletic mode of origin.

Archaeological evidence Carbonized seeds of grass pea appear in several south-western Asian, Aegean, and west Mediterranean Neolithic settlements. To-date, the oldest remains come from ca. 10,250–9,550 cal BP Çayönü, Turkey (van Zeist 1972; van Zeist and de Roller 1991-2, 2003) and ca. 9,450–9,300 cal BP Jarmo, Iraq (Helbaek 1960). A few grass pea seeds were also discovered in Aceramic Neolithic, ca. 8,350–7,750 cal BP, Tepe Sabz, Iran (Helbaek 1970), and Nevali Çori, Turkey (Pasternak 1998). It is impossible to decide whether they represent domesticated forms or collection from the wild. Over 800 charred seeds were found in flotation samples from ca. 8,500–7,500 cal BP Final PPNB Gritille, on the Euphrates, south-east Turkey (Miller 1991, 2002). Such a large quantity is likely to be a crop. The bulk of Neolithic grass pea finds comes, however, from Greece and Bulgaria (Kislev 1989a). About 200 seeds were discovered in ca.8,000 BP Prodromus (Halstead and Jones 1980). A seven-litre hoard was uncovered in ca. 6,500–5,000

cal BP Late Neolithic Servia, where Hubbard (Hubbard 2000), claim by ‘circumstantial evidence’ that toxins were leached by soaking, and some more in Late Neolithic, ca. 6,650–6,300 cal BP, Dimini (Kroll 1979). Numerous seeds were also retrieved from early Neolithic, ca. 7,850–7,500 cal BP, Azmaška Mogila (Hopf 1973a), as well as in contemporaneous storage from Kapitan Dimitrievo, Bulgaria (Marinova 2006, forthcoming). The large stores of seeds found at Late Chalcolithic Kuruçay (3,700– 3,200 cal BC) in south-west Turkey are consistent with an Aegean and south Balkan centre of importance for this crop (Nesbitt 1996). A few Lathyrus-type carbonized seeds were uncovered also in several Impressed Ware and Chasséen sites in south France (Courtin and Erroux 1974). In the Bronze Age, remains of grass pea continue to appear in south-west Asia and in southeastern Europe (for a compilation of data see Kislev 1989b). They were discovered, for example, in Early Bronze Age Lachish, Israel (Helbaek 1958), in Tell Basmosian, Iraq (Helbaek 1963), in ca. 4,500–4,000 cal BP Early Bronze Age Kastanas, Greece (Kroll 1983; see also Fig. 29), and in Middle Bronze Age Tiszaalpár Várdomb, Hungary (Hartyáni 1982). In most of these finds the seeds are small and the seed coats are missing. It is therefore impossible to decide whether they represent a collection from the wild, an infestation of L. cicera-type weeds in fields of other crops, or domesticated forms. Yet the sheer quantities of this pulse in Gritille, Prodromus, Servia, Azmaška Mogila, Kapitan Dimitrievo, and Dimini, seem to indicate domestication. Finally, grass pea is also reported from several Harappa culture sites in Pakistan and north-west India (Fuller 2000). The grass pea might belong to a group of founder crops that started Early Neolithic crop assemblage, or it may have been added soon after the establishment of grain agriculture. When, where, and how this happened is still difficult to say, more so since the wild relatives of this pulse are widely distributed over the entire Mediterranean basin. The early finds from Çayönü and Jarmo, as well the large number in Gritille, suggest that L. sativus may have been taken into domestication in south-west Asia, yet the proposal that it is south Balkan in origin (Kislev 1989b) is another valid option.

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Archaeological evidence Relative to other legumes, finds of L. clymenum are rather rare. The earliest and largest find came from ca. 4,850–4,550 cal BP Early Bronze Age II levels of Yenibademli Höyük on the Gökçeada island, Turkey (Oybak-Dönmez 2005). Seeds of L. clymenum have been discovered in a storage room in ca. 3,900–3,700 cal BP Middle Bronze Age IIA Tel Nami, a coastal site in Israel, suggesting the transport of this pulse from the Aegean basin into the Levant by maritime traders (Kislev 1993). Large quantities of charred seed of L. clymenum, placed also in storage jars, were discovered in Akrotiri, Thera Island, in a house destroyed by the volcanic eruption that devastated this island in ca. 3,578 cal BP (Sarpaki and Jones 1990). These authors also reported seeds of this pulse among plant remains retrieved from Late Minoan II Knossos, Crete, and contemporary Phylakopi, Melos. These finds establishes L. clymenum as a local, Aegean, Bronze Age domestic plant, which survives today only as a relic.

Fenugreek: Trigonella foenum-graecum 0

10 mm

Fig. 29 Carbonized seeds of grass pea, Lathyrus sativus, cotyledons only, no seed coat survived charring, Bronze Age Kastanas, Greece (Kroll 1983).

Spanish vechling: Lathyrus clymenum Lathyrus clymenum L. is a Mediterranean grain crop of restricted distribution, still grown today in several Aegean islands such as Thera (Santorini), Anafi, and Karpathos (Sarpaki and Jones 1990). Like other legumes, the seeds are protein-rich and highly nutritious. L. clymenum is known as a fodder crop, as well as a staple food. Like Lathyrus, the seeds are toxic and might cause lathyrism (Melamed et al. 2009). Wild and weedy forms of L. clymenum are widely distributed in the western and central parts of the Mediterranean basin, from west Turkey to the Iberian Peninsula, and from Cyrenaica to Morocco. Previously, the living plant was unknown in the Levant (Kislev, 1993), but recently Melamed et al. (2009) reported four wild populations in Israel.

Fenugreek, Trigonella foenum-graecum L., (Leguminosae) is a minor pulse crop of Mediterranean agriculture grown in southern Europe, south-west Asia, Ethiopia, and the northern parts of the Indian subcontinent. The seed is widely used as a condiment and as an important flavoring ingredient for the preparation of curry powder and soups. Fenugreek is also used today as an effective nitrogen fixer and effective soil renovator (Small 1997). It is an annual, diploid (2n = 2x = 16 chromosomes), self-pollinated, legume with long curved, linear-lanceolate, stiff pods (mostly growing to a height of 30–60 cm), and characteristic roughly quadrangular seeds which have prominent radicles (‘beaks’). The plant has a long taproot with numerous fine laterals bearing nitrogen-fixing nodules. The crop shows very close morphological resemblance to a group of wild Trigonella species distributed over the east-Mediterranean basin and south-west Asia (Širjaev 1932; Ladizinsky and Vosa 1986; Small 1997). It is not yet clear which

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member of this group (Section Foenum graecum Širjaev) gave rise to the domesticated fenugreek. Yet since all these wild relatives have small distribution areas and are restricted to south-west Asia and the eastern parts of the Mediterranean basin, it seems likely that T. foenum-graecum was brought into domestication somewhere within this geographic region.

Archaeological evidence Charred fenugreek seeds are quite rare in archaeobotanical assemblages. Some were retrieved from south-west Asian sites such as ca. 6,000 BP Tell Halaf, Iraq (Neuweiler 1935), Iron Age Lachish, Israel (Helbaek, 1958), and Iron Age Deir Alla, Jordan (Neef 1989). In addition, desiccated seeds were found in ca. 5,450–5,650 cal BP Ma’adi (Neuweiler 1946), and in several baskets, mixed with coriander, in eighteenth dynasty, ca. 1,325 BC Tutankhamun tomb (Neuweiler 1946; Hepper 1990; de Vartavan et al. 2010) in Egypt. The available archaeological evidence on this pulse is still fragmentary, yet it indicates that fenugreek was part of south-west Asian agriculture already in the Early Bronze Age.

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Lupins: Lupinus Several annual members of the genus Lupinus L., distributed over the Mediterranean basin, have been introduced into domestication and are grown today either as grain crop or as forage plants and green manure. Prominent among them are the white lupin, L. albus L., the yellow lupin, L. luteus L., and the narrow-leaved lupin, L. angustifolius L. (Gladstones 1974; Hanelt 1986; Hill 1995; Cowling et al. 1998). All are vigorous growers and produce large, attractive seed. However, their use is complicated by the fact that lupins generally contain bitter alkaloids which are difficult to remove. Domestication of these grain pulses required an initial selection for non-bitter seed and non-dehiscent pods. The white lupin, L. albus [= L. termis Forssk.], is a traditional Mediterranean pulse crop, which has been grown in this area since antiquity. Even today the white lupin is an appreciated food crop and it is still commonly cultivated in some Mediterranean countries,particularly in Egypt. The wild progenitor of the domesticated white lupin is well identified. The cultivars are inter-fertile and chromosomally identical (2n = 2x = 50 chromosomes) with wild forms, which are native to the southern part of the Balkans, the Aegean islands,

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Map 12 Geographical distribution of wild white lupin, Lupinus albus subsp. graecus [= L. graecus] (based on Gladstones 1974).

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and the Aegean belt in west Turkey (Map 12). They also extend to the Sila Mountains in south Italy (but are very rare there). These were traditionally called L. graecus Boiss. and Spruner. More recently they were taxonomically included in the crop complex as L. albus subsp. graecus (Boiss. and Spruner) Franco and Silva. Wild forms are characterized by dehiscent pods, smaller, more bitter seed, and thicker, impermeable, pigmented seed coats.

Archaeological evidence Earliest L. albus find is reported from ca. 5,450–5,650 cal BP, Late Neolithic, Predynastic Maadi, Egypt, but this single seed cannot serve as evidence for its domestication status and cultivation in Predynastic Egypt (van Zeist and de Roller 2003). Bronze Age remains of white lupin were found in Thera island,

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Greece (Sarpaki 1992b), and a single find is available from Late Bronze Age Hala Sultan Tekke, Cyprus (Hjelmqvist 1979a). Desiccated seed of L. albus—with their characteristic smooth, white coats—were reported from several Roman sites in Egypt (Germer 1985). They were substantiated by additional finds in Qasr Ibrim (Rowley-Conwy 1994), Mons Claudianus (van der Veen 1998, 2001a), and Berenike and Shenshef (Vermeeren and Cappers 2002; Cappers 2006). In conclusion, the distribution of the wild forms of L. albus seems to indicate that this pulse was taken into domestication somewhere in the Aegean basin. When this happened is hard to say since preRoman finds are very scanty. However, the total absence of this crop in Neolithic sites in the Mediterranean basin and Anatolia suggests a late domestication.

C H A PTER 5

Oil- and fibre-producing crops

Indigenous plants producing oil and/or fibre were also taken into cultivation at the start of agriculture in south-west Asia and the Mediterranean basin during the middle-late PPNB, ca. tenth-ninth millennia BP. Oil and fat are an important component of human’s diet, and most societies have a source of fat/oil from plants or animals. Thus, these sources are always an important component of agricultural systems. Flax is apparently the earliest and the best-documented oil and fibre crop. Remains of linseed and fragments of linen indicate that flax belongs to the first wave founder crops of south-west Asia and that it maintained its leading role in the Old World’s fibre and oil production crop from Neolithic times to the early years of the twentieth century. Two other important fibre crops are hemp and cotton. The evidence for the beginnings of their domestication is still insufficient, but it is clear that they were introduced into cultivation outside south-west Asia. However, both hemp (in East Asia) and cotton (in the Indian subcontinent) were already under cultivation in the fourth millennium BP. Several oil plants seem to have entered cultivation not directly, but first by evolving weedy forms. They were added to the crop assembly only after the firm establishment of the principal founder crops such as wheat, barley, lentil, pea, and flax. The case of the gold of pleasure, Camelina sativa (p. 111) is well documented. Several other cruciferous plants (e.g. Sinapis and members of the genus Brassica) seem to have followed a similar mode of evolution under domestication. In a similar way to the cereals and the pulses, the introduction of oil-bearing seed plants into agriculture triggered the evolution of character100

istic domestication traits. Under the new system of sowing, reaping, and threshing, selection brought about the breakdown of the wild mode of seed dispersal and the retention of the seed on the mother plant. Indeed, in most domesticated oil crops the fruits do not dehisce. In others there is a delay in fruit opening. Another conspicuous evolutionary trend is yield increase. Compared with their wild relatives, most cultivars of oilbearing seed crops have frequently larger inflorescences and/or bigger fruits (with larger or more numerous seeds). The wild mode of germination inhibition has also broken down and as a result, seed coats have become thinner. Finally, cultivated varieties frequently show increased oil content. Yet this is not a general rule. Seeds of some wild types are as rich in oil as their cultivated counterparts. In fibre-producing plants the accent is on long, strong fibres. If stem fibres are used (as in flax and hemp), long and straight plants evolve. If lint is the goal (as in cotton), mutations determining long lint (not found in the wild) are selected. We still know very little about how oil and fibres were extracted from plants in the early phases of agriculture. However, it seems reasonable to assume that: (i) Oil could be obtained by decantation; i.e. by crushing the seed, pouring hot water on the meal, and scooping the oil after setting. In some cases, before crushing, the seeds were softened by allowing them to absorb water. (ii) Flax and hemp stems were retted and broken in order to free the fibres. Possibly both technologies antedate agriculture.

OIL- AND FIBRE-PRODUCING CROPS

Flax: Linum usitatissimum Flax, Linum usitatissimum L. (Lineaceae), is an annual, self-pollinated crop with characteristic slender, strong stems and rounded capsules which (in cultivated forms) do not dehisce, but retain the oval, compressed, shining seed. The crop is diploid (2n = 2x = 30 chromosomes) and predominantly self-pollinated. Consequently, in this crop, variation has been moulded in the form of numerous true breeding lines and aggregates of land races. Two specializations are apparent: (i) oil varieties that are relatively short (30–70 cm), branched, and usually bear large seed. They are grown for high yield of linseed; (ii) fibre varieties that are taller, sparsely branched, and usually bear small seed. Transitional forms cultivated for both oil and fibre occur as well. Flax was a principal oil and fibre source in the Old World and probably the earliest cultivated plant used for weaving clothes. Until recently, flax was extensively grown all over the area from the Atlantic coast of Europe in the west, to Russia and India in the east, and Ethiopia in the south (Durrant 1976). In antiquity, flax fibres (which are stronger than cotton or wool) were the principal vegetable fibre used for weaving textiles in Europe and western Asia (Plate 12). However, from the industrial revolution onwards, flax was gradually replaced by cotton imports, and currently it has been almost completely replaced by cotton and synthetic fibres. The seed contains about 40% oil and in peasant communities, linseed was used as a source for edible oil and highgrade lighting oil. Cooking oil is obtained from linseed by cold pressing. Drying oil (used for paints and varnishes) is produced by hot treatment of the seed before pressing (hot pressing). The fibres for spinning are obtained from fibrecell bundles running the length of the stem and forming a ring in the cortex. The stems are harvested before the maturation of the seed. Traditionally they were first dried, and then immersed (‘wetted’) in water to allow the microbial decomposition (‘retting’) of the pectin connecting the fibres with other cells and tissues of the stem. After retting, the stems were dried and the fibres

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(averaging 4 cm in length) were separated by pounding (‘breaking’) and combing. Flax is represented in archaeological excavations both by seed and, occasionally, capsules, and by remnants of stems or textiles. In the latter, fibres can be identified microscopically if they are not carbonized (Körber-Grohne 1991).

Wild ancestry Linum L. is a relatively large genus comprising some 200 species spread over the temperate, Mediterranean, and steppe belts of the northern hemisphere. Domesticated flax, Linum usitatissimum L., is most closely related to wild L. bienne Mill. [= L. angustifolium Huds.]. These two flaxes have the same chromosome number (2n = 2x = 30 chromosomes), inter-cross readily, and are fully inter-fertile (Gill and Yermanos 1967). L. bienne (Fig. 30), with its characteristic strong branches, blue flowers, and dehiscent capsules is widely distributed over west Europe, the Mediterranean basin, North Africa, south-west Asia, and Caucasia (Map 13). Some wild forms are biennial or perennial, others are annual, all are predominantly self-pollinated. L. bienne grows mainly in wet places such as moist grassy areas, springs, seepage areas on rocky slopes, moist clay soils, and marshy lands. Occasionally it is also found as a weed in fields and at the edges of cultivation (including flax cultivation). In such places, hybridization between wild and domesticated flaxes occurs sporadically. On the basis of its close morphological and genetic affinities to the cultivated crop, L. bienne is identified as the wild progenitor of L. usitatissimum (Diederichsen and Hammer 1995). It is therefore correct to regard this wild flax as part of the crop species; i.e. as the wild subspecies of the crop. As in many other grain crops, the main changes under domestication are the shift to non-splitting capsules, the increase of seed size, and the selection for higher oil yield, or longer stems with a high amount of long fibres. Fu (2005) studied 2727 worldwide accessions of flax using RAPD markers, and Allaby et al. (2005) have studied the sad2 locus in both wild and domesticated accessions of flax. In addition, Fu and Allaby (2010) assessed phylogenetic relationships among twenty-nine Linum accessions from sixteen wild

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5 cm

0

A

B

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Fig. 30 Wild flax, Linum usitatissimum subsp. bienne [= L. bienne]. A–Flowering and fruiting stem; B–Capsule (Zohary 1972, plate 374).

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Map 13 Geographical distribution of wild flax, Linum usitatissimum subsp. bienne [= L. bienne].

and domesticated species, based on four non-coding regions of chloroplast DNA sequences. The differences in genetic diversity between and within accessions revealed in these studies suggest that domesticated flax is of a monophyletic origin, it has probably evolved from a single domesticating event, although the geographic location of this event is still obscure. In addition, these data indicate that flax domestication selected for the larger, oil-rich, seeds of the oil producing variety, rather than its fibers (Allaby et al. 2005; Fu and Allaby 2010). In other words, these studies suggest that flax was first domesticated as an oil plant.

Archaeological evidence Flax was apparently used by humans already before its domestication. Archaeological finds of seeds are more likely to result from cultivation (or gathering) for seed consumption (as McCorrison 1997), since harvesting for fibers often occurs before seed maturation. Recently, twisted and dyed flax fibres were reported in Upper Palaeolithic, ca. 30,000 year-old, Dzudzuana Cave, Georgia (Kvavadze et al. 2009,

but see discussion by Bergfjord et al. 2010; Kvavadze et al. 2010). The oldest wild linseed remains retrieved from archaeological sites in south-west Asia come from ca. 11,800–11,300 cal BP Tell Mureybit (van Zeist and Casparie 1968; van Zeist and BakkerHeeres 1986). Soon after, seeds of flax were found in many of the Pre-Pottery Neolithic B farming villages that appeared in the Fertile Crescent from 10,500 cal BP onward (Map 1). Some of the earlier finds come from Çayönü, Turkey (van Zeist 1972; van Zeist and de Roller 1991-2, 2003), Tell Aswad, near Damascus, Syria (van Zeist and Bakker-Heeres 1985), Ali Kosh, Iran, (Helbaek 1969), Jericho, Israel (Hopf 1983), and ‘Ain Ghazal, Jordan (Rollefson et al. 1985). The seeds are still small, similar in size to those of wild bienne forms. Yet they are almost always associated with domesticated wheats and barley. Fragments of a capsule from 9,900–9,550 cal BP Middle PPNB Jericho (Hopf 1983, Fig. 31) are probably the earliest indication we have today of domesticated flax. Another indication of early flax domestication comes from linseed remains recovered from Late PPNB, 8,700–8,600 cal BP, levels of

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Tell Ramad, Syria (Fig. 32). The calculated size of the seeds from this site, corrected for charring shrinkage, ranges from 3.2 to 4.1 mm in length. This is already within the size class of the L. usitatissimum seed, the lower limit of which lies at 3.0 mm. It is therefore an attractive indication for flax domestication under rain-dependent conditions before ca. 8,600 cal BP (van Zeist and Bakker-Heeres 1975). Such domesticated linseeds were found in Pottery Neolithic Nahal Zehora, Mount Carmel, Israel, ca. 8,150–7,850 cal BP (Kislev and Hartmann forthcoming). In the Mesopotamian basin, linseed sizes in ca. 8,350–7,750 cal BP Tell Sabz, Iran (Helbaek 1969) and in Halafian, ca. 7,750–7,250 cal BP Arpachiya (Helbaek 1959a, 1959b) are even bigger (4.7–4.8 mm long). As argued by Helbaek, these large seeds indicate advanced domestication and demonstrate that flax was part of the irrigated grain agriculture system that evolved in this region. In Choga Mami, Iraq, Helbaek (1972) reports a rare find in the earlier, ca. 7,800–7,750 cal BP, stratum, which became frequent at the following stratum, dated to the second half of eighth millennium. Linseed also appears in several later south-west Asian Neolithic and Bronze Age sites. A complete capsule was retrieved from Early Bronze Age, ca. 3,500–3,300 cal BP, Jericho (Fig. 31). Flax is associated with the spread of Neolithic agriculture from the Fertile Crescent ‘core area’ into Europe and the Nile Valley (Map 2—Plate 6) from the early Neolithic (eighth millennium BP) onward. Linseed has been recovered from sites dating from the first half of the eighth millennium BP in the

0

4

Mediterranean Basin and southern parts of Europe. Such sites are Knossos, Crete (Sarpaki 2009), a few sites in Thessaly, Greece (Kroll 1981b, 1991), La Marmotta, Italy (Rottoli 1993, 2002), and as far as Mohelnice, Moravia (Opravil 1979, 1981; Kühn 1981). Further north, in central and western Europe, the earliest cultivated flax finds are from the second half of the eighth millennium BP. They belong to the earliest Linearbandkeramik, food production culture in the area. Sites include those in the Aldenhovener Platte (Knörzer 1973, 1974, 1997) and Rosdorf (Kirleis and Willerding 2008) in Germany, Brześć in Poland (Bieniek 2007), and North Meseta sites (including Cueva de La Vaquera, La Lámpara, and Revilla del Campo) (López et al. 2003; Stika 2005; Peña-Chocarro 2007), Spain. The earliest find in Austria, however came from Late Linearbandkeramik, ca. 7,150–6,800 cal BP, Schletz (Schneider 1994; Kohler-Schneider 2007). In Switzerland it occurs among plant remains in many lake-shore settlements (Jacomet 2007) but arrives only in the second half of the seventh millennium BP. It is still rare in the relatively early (ca. 6,250 cal BP) Egolzwil 3 culture (Bollinger 1994; Jacomet 2007), but becomes frequent in the later Neolithic and Bronze Age times, in the settlements at Lake Zürich or at Late Neolithic Lake Biel (Ammann et al. 1981; Brombacher 1997, 2000; Brombacher and Jacomet 2003). At the second half of the sixth millennium BP, a few seeds, capsules, and textile were found in Station III de Clairvaux, France (LundströmBaudais 1984). At Late Bronze Age, 2,905–2,869

8 mm

Fig. 31 Carbonized capsule of domesticated flax, Linum usitatissimum subsp. usitatissimum, Bronze Age Jericho (Hopf 1983).

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Ramad

Ras Shamra

0

3 mm

Fig. 32 Carbonized seeds of domesticated flax, Linum usitatissimum subsp. usitatissimum, from Neolithic Ramad, and Ras Shamra, Syria (van Zeist and Bakker-Heeres 1975).

(dendrodated) BP, Grésine, France, linseed and capsules were frequently found (Bouby and Billaud 2001). Flax is similarly common in lake-shore settlements in fifth millennium BP south Germany, such as Ödenahlen at the Federseeried (Maier 1995). At the same time, linseed was found for the first time in Late Neolithic Aartswoud, The Netherlands (Pals 1984). Linseed was also retrieved from late Neolithic and Early Bronze Age northern Italy; that is, from Lagozza (Buschan 1895), from Valeggio (Villaretvon Rochow 1958), and Frioli (Pessina and Rottoli 1996). In Rumania, lumps of charred linseed were discovered in Sucidava-Celei (Cârciumaru 1996), in contexts dated to the transition between the Eneolithic and the Bronze Age (ca. 4,850–4,150 cal BP). The earliest linseed, including fibers, in Spain and Portugal were uncovered in Chalcolithic (ca. 4,800–4,200 cal BP) Buraco da Pala, Portugal and several El Argar culture, ca. 4,150–3,350 cal BP, Early Bronze Age sites in Spain. The earliest documentation of flax cultivation in Scandinavia appears in Bronze Age, ca. 3,150–2,950 cal BP, Borge vestre, Norway (Sandvik 2007, 2008), while in Denmark strong indication for flax agriculture and industry revealed first in Late Bonze Age, ca. 2,800 cal BP, Frydenlund (Henriksen and Runge 2009).

In Egypt, flax was retrieved (together with emmer wheat and barley) in the earliest Neolithic farming sites discovered in the Nile Valley during the eighth millennium BP, Fayum (Caton-Thompson and Gardner 1934; Wetterstrom 1993; Wendrich and Cappers 2005) and Merimde (Wetterstrom 1993, 1998). Remains of flax seeds were also found in ca. 5,200 BP El Omari (Wetterstrom 1993), and they continue to appear in several other pre-dynastic sites. From the start of dynastic times onwards, flax emerges as one of the founder principal grain crops of Egypt. It kept this role until very recently. Remains of flax textiles also make an early appearance, attesting that fiber varieties developed not much later than the oil-producing varieties. The best examples come from the drier parts of southwest Asia, where due to low humidity, woven material survived without carbonization. Pieces of exquisitely woven linen were discovered among Pre-Pottery Neolithic B (beginning of the ninth millennium BP) remains in Nahal Hemar Cave near the southern tip of Dead Sea, Israel (Schick 1988). A single piece of linen was found in Neolithic (eighth millennium BP) Fayum, Egypt (Caton-Thompson and Gardner 1934). Linen fragments were retrieved from the Chalcolithic, ca. 6,200–6,000 cal BP, ‘Cave of the Treasure’ near the Dead Sea, Israel (Bar-Adon

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1980). The most remarkable linen find was discovered in contemporaneous ‘Cave of the Warrior’ near Jericho. It includes a beautifully preserved, colored, and fringed kilt and sash, and large (seven meters long and two meters wide) shroud, wrapping the body of a dead nobleman. These finds indicate a high level of familiarity and expertise in domesticated linen production, already extant in this period (Schick 1998, 2003). Flax textiles were common in the Old, Middle, and New Kingdoms of Egypt, where linen was extensively used for wrapping mummies (Täckholm 1976; Germer 1985) Egyptian retting, spinning, and weaving of flax is beautifully recorded in the twelfth-dynasty Beni Hasan grave paintings (Täckholm 1976). In summary, the available archaeological evidence clearly suggests that flax belongs to the first group of grain crops that started agriculture in the Levant during the Middle PPNB. As for distinguishing the domesticated from the wild flax, the situation here is parallel to the cereals and pulses; it is manifested in the function of the seed-dispersal apparatus (diaspore). In wild populations, the diaspores split and disperse the seeds immediately after maturation. In contrast, breakage of the wild type seed-dispersal mechanism does not occur in the domesticated type and the capsule stays intact and waits for the reaper. Hence, separated seeddispersal units in the archaeobotanical assemblage indicate the wild type, while whole capsule, or fused diaspores, indicate domesticated type. In addition, the gradual increase in seed size and the use of linen indicate that flax domestication was, very probably, already practised in the PPNB Levant, as attested first by Middle PPNB capsules and then by Late PPNB seeds and linen textile. The evidence from the living plants shows that wild bienne forms are widespread in the ‘arc’, and fully support the contention that domestication occurred in the Fertile Crescent. Molecular studies also suggest single domesticated events for flax, of the oilproducing variety.

Hemp: Cannabis sativa Hemp, Cannabis sativa L., is a tall (2–3.6 m) dioecious, wind-pollinated herb, and a member of a special small family Cannabaceae. Hemp is an

extraordinarily variable taxon that contains wild forms, weedy types, and domestic cultivars. It is grown for three main purposes: (i) for fibres obtained from the bast of the stems; (ii) for the seeds that are used either for extraction of oil or as an animal feed; and (iii) as a source of a psychoactive drug produced by the glandular hairs of the plant. Special cultivar groups have been developed for the different uses (Small 1995). Fibre varieties are tall, succeed in both temperate and tropical climates, and contain negligible quantities of the drug. Hemp textiles are strong but coarse and in traditional communities they were used for cheap clothing, sacks, rags, or sails. Hemp oil is used mainly for technical purposes such as varnish or soap. The drug is obtained from special varieties (frequently referred to as the indica group) grown in warm climates. Only the tops of the female plants contain appreciable quantities of the Cannabis drug and are harvested for the preparation of marijuana (or hashish). Most taxonomists consider C. sativa and C. indica to be one species, but Hillig (2005), based on allozyme variation at 17 gene loci from 157 Cannabis accessions, suggest that Cannabis has derived from two major gene pools. On the basis of this data, he recognized C. sativa and C. indica as separate species, both with seeds and fibre landraces. Domesticated hemp is closely related to, and fully inter-fertile with, an extraordinarily variable aggregate of wild and weedy forms. Except for traits associated with the economic use, wild forms differ from the cultivars by relatively smaller achenes (‘seed’), which also possess adhering perianths and elongated bases. The crop is mostly a central Asiatic element. Temperate territories in this vast area such as the Caspian Basin, parts of Afghanistan, central Asia, and the Himalayas, harbour spontaneous C. sativa plants that seem to be wild. Very likely, populations that appear to grow wild are not fully indigenous, but have been introgressed extensively with the cultivated varieties and weedy types which also abound in the same areas (Small and Cronquist 1976). The picture of Cannabis is even further complicated by the marked tendency of this cross-pollinated crop to revert to wild. Naturalized derivatives and weedy races of C. sativa are now distributed not only in central Asia, but also all over the world.

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Archaeological records on the early establishment of hemp are not available so far. However, this plant must have been taken into domestication quite early, somewhere in temperate Asia. On the basis of linguistic and cultural evidence Li (1974) concluded that hemp was probably grown in China by at least 4500 BP and that it was the only fibre available to the ancient peoples of northern and north-eastern China. Merlin (2003) summarizes the available evidence, archaeological and otherwise, of hemp, as well as other psychoactive plants in the old world. The recent expansion of hemp to south-west Asia, the Mediterranean basin and Europe is now well documented. Remains of hemp fabrics are available from ca. eighth century BC Gordion, Anatolia (Bellinger 1962). It also seems certain that the plant was known to the Sarmatians and Scythians who occupied the southern part of Russia between approximately 700 and 300 BC (Godwin 1967). Herodotus (ca. 446 BC), describes a Scythians purification ritual: ‘On a framework of three sticks meeting at the top, they stretch pieces of woolen cloth, taking care to get the joins as perfect as they can, and inside this little tent they put a dish with redhot stones on it. Then they take some hemp seed, creep into the tent, and throw the seeds on the hot stones. At once it began to smoke, giving off vapor unsurpassed by any vapor-bath one could find in Greece. The Scythians enjoy it so much that they would howl with pleasure’ (Herodotus, IV, 75, after Merlin 2003). Such a practice is attested by a special find in a series of fifth-century BC frozen kurgans (burial mounds) at Pazyryk, Siberia. A 1.2 m-high wooden frame tent was found within each burial, and a bronze vessel inside it. Stones and hemp seeds were found inside each vessel. ‘A leather pouch with hemp seeds provided supplies, and scattered hemp, coriander and melilot seeds were also recovered’ (Rudenko 1970, as cited by Merlin 2003; Sherratt 1995). Despite the appeal of such a find, hemp seeds can be confused with those of Panicum miliaceum (Bakels 2003), for example, re-identification is therefore advised. At contemporaneous sites in Ukraine, Pashkevich (1999) found Cannabis seeds in several Scythian sites. At that time hemp became a well-known fibre crop in south-west Asia and Greece. It spread to

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Italy and Sicily in about 100 BC. The fibres were used, among other things, to make ropes and sails. An interesting find of both fruits and fibres came from the Iron Age, ca. 2,400–2,050 cal BP, salt-mining site and settlement Dürrnberg/Hallein, Austria (Werneck 1949; Swidrak 1999; Boenke 2007). The spread of C. sativa to central and north Europe from Roman times on is now documented by a wealth of macro-remains, palynological evidence, and written sources (Dörfler 1990). The narcotic properties of C. sativa were recognized in India by ca. 1000 BC.

Old World cottons: Gossypium arboreum and G. herbaceum Gossypium is a large genus of the family Malvaceae containing some fifty species, distributed over large landmasses in tropical and subtropical warm environments. There are four different cotton crops, two species in the Old World (G. arboreum L. and G. herbaceum L.), and another two in the New World (G. hirsutum L. and G. barbadense L.). An important difference between the domestic species of the Old World and the New World is that Old World cotton crops are all diploid (2n = 2x = 26 chromosomes, genomic designation AA), whereas the New World crops are tetraploid (2n = 4x = 52 chromosomes, genomic designation AADD). In this book we are dealing with the Old World material only. There is a revival of cotton as one of the most important commercial plants for the present-day textile industry. Both G. herbaceum and G. arboreum bear spinnable but relatively short (less than 22 mm long) seed fibres or lint (Lee 1984). These were the only Gossypium crops grown in the warmer parts of Asia and Africa before the discovery of America. According to Fryxell (1984, p. 30) the widest crop diversity of G. arboreum was in the Indian subcontinent, while the center of G. herbaceum’s diversity was in East Africa and in the Levant. In post-Columbus times, Old World cottons have been almost fully replaced by their American counterparts (G. hirsutum and G. barbadense), which produce longer (2434 mm long) lint. Today G. arboreum and G. herbaceum survive as relic crops, and account for less than 1% of the world cotton production. For a survey of both domesticated and wild cottons consult Lee (1984),

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Fryxell (1979, 1984), Wendel (1995), and Wendel and Cronn (2003). Butterworth et al. (2009) described recently the morphological transformation undergone by cotton fibers in the course of domestication; from short, coarse fibers in the progenitor wild species, to the long, fine fibers in the domesticated species. Both G. arboreum and G. herbaceum are plants growing in warm climates, and basically are perennial crops (the first can reach the size of a small tree). However, under domestication some annual cultivars evolved as well, in both crops, and rendered cotton cultivation possible in areas with relatively cold winters. The two species, G. arboreum and G. herbaceum, are obviously closely related genetically. They are similar morphologically, cross readily, and their F1 hybrids are almost fully fertile. Both carry homologous chromosomes (genome A) and an identical chloroplast genome. When grown side by side, they tend to hybridize spontaneously. In such cases, their morphological boundaries are frequently blurred. This suggests that both cultigens belong to the same crop species complex and could have evolved from a single wild ancestral stock. On the other hand, domestic G. arboreum and G. herbaceum were found to differ from one another by a chromosomal translocation, and to manifest hybrid breakdown at the second (F2) and the third (F3) hybrid generations (Phillips 1961). Moreover, isozyme tests showed that the two crops are characterized by numerous distinct alleles, indicating that the genetic similarity between them is lower than the values known to occur in most crop-ancestor species pairs (Wendel et al. 1989, 2003). As argued by these authors, such divergence does not support the notion of two cultigens derived from a common wild ancestor. Instead, it suggests that each evolved from a distinct wild stock. Moreover, they indicate that cotton is unique among crop plants in that four separate species (the two Old World species and the two New World species) were independently domesticated. The present knowledge of the close wild relatives of the Old World cultivated cottons (wild diploid taxa containing genome A) is still insufficient, and of little help in solving the problems of how and where G. arboreum and G. herbaceum were domesticated. Wild forms of G. herbaceum have already been discovered and identified both taxonomically and

cytogenetically. They are referred to today as G. herbaceum L. subsp. africanum (Watt) Hutch. These wild taxa are known to grow only in southern Africa (Vollesen 1987), a region where agriculture arrived at a very late date, and geographically a place widely separated from the traditional territory of G. herbaceum cultivation. The situation in G. arboreum is even more perplexing. Gossypium taxonomists (Fryxell 1979, 1984; Vollensen 1987) do not mention any wild arboreum forms in their studies. It is not clear whether this is a real absence or only a reflection of the paucity of field findings. All in all, wild diploid cottons—containing genome A—have not yet been conclusively detected in the geographic areas in which they should be expected—Africa north of the equator, south Arabia and/or the Indian subcontinent.

Archaeological evidence The archaeobotanical documentation of cotton is far from being satisfactory. The earliest evidence of cotton fibers came from ca. 8,000–6,500 BP Ceramic Neolithic Mehrgarh, Baluchistan (Moulherat et al. 2002). The fibers were identified as Gossypium sp., most probably represent wild origin. Later, reliable signs of cotton use, as attested by fibers and seeds finds, come from Harappan sites (ca. 4,250–3,750 BP) in the Indian subcontinent. Here fragments of cotton textiles and strings, preserved by copper and silver oxides, were uncovered in Mohenjo-Daro, Pakistan (Gullati and Turner 1929). In addition, cotton remains are reported from contemporary Harappa and several other Indian and Pakistani sites (Fuller 2000, 2008). As stressed by Meadow (1996), these finds indicate that Gossypium fibers were exploited in the Indus valley as early as the end of the fifth millennium BP. However, it is unclear whether these early remains represent arboreum or herbaceum cottons. A more problematic early find comes from ca. 4,500 BP Afyea, Egyptian Nubia. Cotton seed and lint hairs (intermediate between those borne by wild forms and those produced by herbaceum cultivars) were discovered in goat coprolites (Chowdhury and Buth 1970, 1971). It is hard to imagine that the Afyea find represents domestic cotton, since it is not accompanied by any other contemporary signs of cotton cultivation or cotton use in the Nile Valley,

OIL- AND FIBRE-PRODUCING CROPS

which is known so well archaeo-botanically. As Moulherat et al. (2002) and Fuller (2008) noted, if the identification of sixth Millennium BP cotton fibers from Dhuweila, Jordan (Betts et al. 1994) will be confirmed, it was most likely imported, perhaps from the Indian subcontinent. Cotton seems to have moved into south-west Asia and the Mediterranean basin fairly late. The earliest sign comes from Assyria where we are informed that ‘trees bearing wool’ were grown by Sennacherib at about 694 BC (Thompson 1949). There are no indications that this early introduction succeeded, but the mentioning of a tree suggests that G. arboreum was involved. Cotton is already referred to in Greek, Roman, and Jewish literary sources, from Hellenistic times onwards. Watson (1983, pp. 32, 34) surveyed the literary information and concluded that already in Greek and Roman times, Indian cotton goods were imported into south-west Asia and the Mediterranean basin, and that they were rare and expensive commodities. The diffusion of cotton cultivation (and of its processing technologies) is another matter. There is no sound literary support for cotton cultivation in south-west Asia at that time. The Jewish sources Mishna and Talmud (particularly the late ones dating to the AD fifth and sixth centuries) refer to several traits of the cotton plant itself, indicating that cotton might already have been grown in the Levant countries (Amar 2000). By compiling current data, Wild (1997) and Decker (2009) demonstrated that cotton cultivation was widespread in Roman Egypt. To-date, the earliest unequivocal archaeobotanical evidence on cotton cultivation outside the Indian subcontinent, comes from late Sassanian (AD sixth and early seventh centuries) Merv in Turkmenistan (Nesbitt 1993, 1994). Fully developed cotton cultivation and sophisticated cotton industry appear in the Levant and the Mediterranean basin only in early Islamic times (Watson 1983; Amar 2000).

Poppy: Papaver somniferum Poppy, Papaver somniferum L. (Papaveraceae), is grown for two purposes (Duke 1973). First, it is famous as a source of opium, which is obtained from the latex released by the plants after gashing their unripe capsules. It is also cultivated for its tasty seeds, which are rich in oil. The seeds are con-

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sumed as such or used to extract poppy oil. Because of this dual use, two series of cultivars have evolved in P. somniferum. The opium forms are grouped in subsp. somniferum Corb., while the oil varieties are collectively known as subsp. hortensis (Hussenot) Corb. The oil is used both for eating and for industrial purposes. The opium is a powerful medicinal and narcotic element. Its effects, including pain killing, were already appreciated in antiquity (Merlin 2003). Domesticated poppy, Papaver somniferum, is predominantly self-pollinated. Most cultivars are diploid (2n = 2x = 22 chromosomes), while wild setigerum DC poppies comprise both diploid and tetraploid (2n = 4x = 44 chromosomes) types. The crop is closely related to wild and weedy forms of poppies that grow, mainly in coastal areas, in the western part of the Mediterranean basin, including most of the islands (Map 14). The diploid setigerum forms were found to be fully inter-fertile with the somniferum cultivars (Hammer and Fritsch 1977). They are therefore assumed to be the progenitor stock from which the cultivated poppy evolved. This necessitated a taxonomic reconsidering of the rank of P. setigerum, which is placed as the wild race, subsp. setigerum (DC.) Corb., of the P. somniferum crop complex (Kadereit 1986). The most conspicuous differences between the tame and the wild plants are the considerable increase in capsule size, and the retention of the seed in the capsule in the poppies under domestication—in domesticated forms the pores beneath the stigmatic lobes do not open. Domestic forms also show a tendency to increased seed size. However, some wild populations bear fairly large seeds which considerably overlap the dimensions (and seed-coat reticulate sculpture, see Fig. 33) found in the cultivated poppy. Seed size is therefore an unreliable trait for recognition of domestication in this crop (Fritsch 1979); which necessitates capsule remains for properly identified domestic form in archaeological assemblages.

Archaeological evidence The earliest wild poppy seed came from ca. 8,000– 7,500 cal BP PPNC Atlit Yam, Carmel Coast, Israel (Kislev et al. 2004). This find might indicate that its distribution area (see Map 14) included Israel dur-

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200 miles 400 km

Map 14 Geographical distribution of wild poppy, Papaver somniferum subsp. setigerum [= P. setigerum] (based on La Valva et al. 1985; Kadereit 1986). Setigerum forms extend westwards beyond the boundaries of this map into Madeira, the Canary, and the Azores Islands.

ing the early Neolithic period. Abundant waterlogged seeds of domesticated poppy came from ca.

Fig. 33 The reticulate sculpture of the seed coat in poppy, Papaver somniferum (Schoch et al. 1988).

7,750–7,150 cal BP Early Neolithic La Marmotta, Italy (Rottoli 1993, 2002). A few carbonized domesticated poppy seeds were found in the Linearbandkeramik settlements of the Aldenhovener Platte, north-western Germany (Knörzer 1971, 1997), Meindling, south Germany (Bakels 1996), and a few more in Zeslawice, Poland (Giżbert 1960), and north of the Alps (Kreuz 2007). Numerous remains of charred seed, as well as occasional capsules, are available from middle and late Neolithic central Europe (for review see Schultze-Motel 1979a; Merlin 2003). They were also retrieved from middle Neolithic Menneville, north France (Bakels 1984), and became common in Late Neolithic (fifth millennium BP) and in Bronze Age lake-shore settlements in Switzerland (Jacomet et al. 1989; Brombacher 1997), in south Germany (Maier 1995; Rösch 1998), and in Lagozza, north Italy (Buschan 1895). Four desiccated, beautifully preserved, wild poppy capsules were reported by Neuweiler (1935) from Cueva de los Murciélagos

OIL- AND FIBRE-PRODUCING CROPS

near Albuñol, Granada, south Spain (despite the same name, this is a different cave from that of Córdoba mentioned above). Significantly, poppies were not discovered in Neolithic sites in southeastern Europe and in southwest Asia. They appear, however, in several Bronze Age sites in Greece, Bulgaria, and the former Yugoslavia (Kroll 1991). As stressed by van Zeist (1980) and by Bakels (1982) this fact, combined with the distribution area of wild setigerum poppies (Map 14), suggests a west Mediterranean domestication. In other words, P. somniferum does not belong to the primary ‘first circle’ Fertile Crescent crops that started food production in Europe. It is a representative of the ‘second circle’ domesticants, (i.e. crops that were added to the original assemblage outside the Fertile Crescent core area).

Gold of pleasure: Camelina sativa Gold of pleasure or false flax, Camelina sativa (L.) Crantz., of the mustard family Cruciferae (Brassicaceae) is a relic oil plant rapidly disappearing from cultivation. Yet until the 1940s, Camelina was an important oil crop in eastern and central Europe. Camelina seed, with their characteristic protruding embryos, appear repeatedly in European archaeological contexts. The crop, subsp. sativa (Mill.) E. Schmid, is closely related to, and inter-fertile with, a variable aggregate of wild and weedy forms distributed over Europe and south-west Asia (for review, see Markgraf 1975). Truly wild, late-flowering forms, now recognized as subsp. microcarpa (Andrz.) E. Schmid, but formerly referred to as C. microcarpa Andrz., grow in east European steppes and adjacent Asiatic territories. They have characteristic small fruits. Closely related to them are early flowering hairy forms, known as subsp. pilosa (DC.) E. Schmid, which thrive in fields of winter cereals over most of Europe. These are obviously recently evolved weeds, which spread from east Europe westward, over cultivated lands. An additional distinct Camelina weedy race, namely subsp. alyssum (Mill.) E. Schmid [= C. sativa var. linicola Prusch.] evolved in association with flax cultivation in Europe. These linicola forms, with their long erect stems and hard fruits, serve as well-documented examples of evo-

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lution of weed mimicry to a specific cultivated crop. The cultivated varieties of C. sativa differ from the wild and weedy forms by their non-dehiscent, large, and pyriform fruits, and by bigger (1.5–2.0 mm long) seed, which contain an appreciable amount (27–31%) of edible oil.

Archaeological evidence Camelina seeds (as well as occasional pods) appear repeatedly in central and eastern European archaeological contexts. However, they start late and the earliest finds come from fifth and fourth millennia BP contexts. Large samples, as well as pure samples; i.e. material that can be confidently regarded as confident domestication appear even later. In central Europe the oldest remains come from final Neolithic contexts (ca 4,000 BC) of Auvernier, Switzerland, and these are followed by several Bronze Age (1,800–1,200 BC) finds from Poland, Hungary, Germany, and north Italy (Schultze-Motel 1979b; Wasylikowa et al. 2001). Iron Age finds are more numerous. Camelina remains become particularly common in the coastal areas of the Baltic and the North Sea. At least some of these finds (particularly the large samples with bigger seeds) seem to represent Camelina cultivars. Further south-east, a single seed of Camelina was found in Chalcolithic (ca. 5,000 BP) Pefkakia, Thessaly (Kroll 1991), and many more in ca. 4,200 BP Sucidava-Celei, Rumania (Cârciumaru 1996). Camelina seeds were also retrieved as a weed of flax from Late Chalcolithic Kuruçay in south-west Anatolia (Nesbitt 1996), from Early Bronze Age Demircihüyük in north-west Anatolia (Schlichtherle 1977/1978), and in Iron Age Yoncatepe in Van Lake province (Dönmez and Belli 2007). They reappear in several Late Bronze Age sites in Greece, Bulgaria, and the former Yugoslavia (Kroll 1991), and also in Late Bronze Age Hadidi on the Syrian-Turkish border (Miller 1991). In the latter site their concentration again suggests cultivation. The combined evidence from the living plants and from archaeology indicates that C. sativa is a secondary crop. Probably this crucifer entered agriculture first by evolving weedy races that infested flax and cereal cultivation. Only later was the weed picked up as an oil crop.

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Other cruciferous oil crops The cultivation of several other members of the mustard family (Cruciferae/Brassicaceae) is amply recorded in classical times. They have been appreciated as oil or mustard sources (extracted from their seed), and/or for their vegetable parts (see Chapter 7). Foremost among these crucifers are: radish Raphanus sativus L., turnip Brassica rapa L. [= B. campestris L.], rape or swede B. napus L. (tetraploid, 2n = 4x = 38 chromosomes), and the various mustards; i.e. the white mustard Sinapis alba L., (diploid, 2n = 2x = 16 chromosomes), black mustard B. nigra (L.) Koch, (tetraploid, 2n = 4x = 36 chromosomes), and brown mustard B. juncea (L.) Czern. (diploid, 2n = 2x = 2 chromosomes). All these crops have wild forms distributed over west Asia and Europe. They include variable, aggressive races of weeds, which infest agricultural land far beyond these territories. It is very likely that seed of wild forms (Brassica, Sinapis, and also Camelina and Descurainia) has been collected or even grown as oil sources in Switzerland and Germany in the late Neolithic times (Schlichtherle 1981). These crucifers were already well-established oil crops in Hellenistic and Roman times. Therefore, it is safe to assume that they were taken into cultivation earlier. Yet, there are almost no archaeological records available for any of these crops. (For the scant information present, see Renfrew 1973, pp. 166–167, and the annual reviews of Schultze-Motel 1968–1994, and Kroll 1995–2000, as well as the online databases http://www.cuminum.de/ archaeobotany/database/; http://www.archaeobotany.de/database.html). Suggestions as to the origin of these plants are necessarily based on linguistic considerations. One rich find of cruciferous seed is described from ca. 5,000 BP Khafadje, Iraq. Here, numerous carbonized seeds were discovered in several locations at the Temple Oval (Bedigian and Harlan 1986). They were studied by E. Schiemann (1933) who concluded that they belonged to Brassica or Sinapis. It is very difficult to distinguish between the seeds of these two genera in charred material. Two cakes with crushed seed of Brassica/Sinapis were found by Willcox (2002) in tenth millenium BP PPNA Jerf el Ahmar, Syria. The presence of Brassica

rapa remains in several late Neolithic and Bronze Age lake-shore sites in Switzerland (e.g. Seeberg, Villaret-von Rochow 1967), suggests a similar use. In other words, turnip may already have been an oil-bearing ‘tolerated weed’ at this relatively early time.

Sesame: Sesamum indicum Sesame, Sesamum indicum L. [= S. orientale L.] (Pedaliaceae), is a traditional, warm-season, annual oil crop whose genus contains some two wild species. Cultivars of this crop have been grown in tropical and subtropical environments, such as south-west Asia and the Mediterranean basin. Sesamum is highly appreciated for its oil that keeps fresh for a long time without turning rancid (Nayar 1995). It was extensively cultivated in the GraecoRoman World (Lenz 1859) both for its attractive oil and for its edible seed. Theophrastus categorizes sesame, along with the millets, as one of the main summer crops of his time. Yet despite such wide use in Roman times, sesame is not an indigenous crop in the Mediterranean and south-west Asian agriculture. The genus Sesamum includes tropical and subtropical wild species, grouped in four sections (Ihlenfeldt and Grabow-Seidensticker 1979). The majority of these species occur in Africa. A few occur in the Indian peninsula and in south-east Asia. All these taxa (except for the crop varieties) do not occur in south-west Asia or in the Mediterranean basin. In fact, this genus is geographically restricted to tropical and subtropical Africa south of the Sahara (the majority of the wild species), and only a few species in the Indian subcontinent and the Far East. A group of wild-growing and weedy forms occur in northwest India, as reported by Bedigian and Harlan (1986) and Bedigian (1998, 2004, 2010) as S. orientale L. var. malabaricum Nar. A second similar wild group was discovered on the southern and western shores of the Indian peninsula, and named S. mulayanum Nair (Hiremath and Patil 1999). Both groups show close morphological and cytogenetic affinities with the crop. Very likely they are elements of the wild stock from which cultivated sesame could have been derived.

OIL- AND FIBRE-PRODUCING CROPS

Archaeological evidence Sesame seeds are very fragile when charred (even more so than other oil-rich seeds). In part, this explains their scarcity in the archaeobotanical record. The oldest record of S. indicum cultivation comes from ca 4,250–3,750 BP Harappa in the Indus Valley (for review see Bedigian 1985, 1998, 2010; Fuller 2003). This was augmented by the find of a small quantity of well-preserved, charred sesame seed in Miri Qalat and Shahi Tump, Markan region, Pakistan, in contexts dated to the second half of the fifth millennium BP (Tengberg 1999). The earliest definite signs of sesame seeds in southwest Asia come from ca. middle of the fifth millennium BP, Early Bronze Age Abu Salabikh, Mesopotamia (Charles 1993). Somewhat later, it appears in ca. 3,200 BP, Late Bronze Age, Tell Sabi Abyad, northern Syria (van Zeist 1994). At adjacent Tell Schech Hamad, van Zeist (2001, 2003) reported sesame seeds from two contexts, a single seed from ca thirty-third century BP and a few more seeds from ca. twenty-seventh century BP. Also in the same period, baskets of sesame seeds were found in Tutankhamun’s tomb in Egypt, ca. 1,325 BC (de Vartavan et al. 2010) The unique circumstances of King Tut perfectly-preserved un-charred funerary offerings most probably represent a rare import, fit for the king’s table—certainly not a local cultivation.

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They are followed by four large finds of Iron Age jars containing carbonized sesame seed that were excavated in Karmir-blur on the outskirts of Yerevan, Armenia, dated to 900 and 600 BC. The site also contains elaborate installations for the extraction of oil from the seed (Bedigian 1985). Another find of the same period comes from the Urartu Kingdom of Bastam, north-west Iran (Hopf and Willerding 1989). In addition, some 200 sesame seeds were uncovered in Iron Age, ca. 800 BC, Deir Alla, Jordan (Neef 1989). It seems that S. indicum arrived to south-west Asia and the Mediterranean basin from the east in the Early Bronze Age (Bennett and Maxted 1997). The relative small number of finds probably indicates small-scale cultivation for culinary purposes rather than large-scale oil production (Neef 1989). It was apparently taken into cultivation in the Indian subcontinent. It is still undecided when and where sesame was domesticated. However, Fuller (2003) and Bedigian (2004, 2010) felt that the combined evidence obtained both from the living plants and from archaeological sites strongly supports the notion that domestication of sesame started in the Indian sub-continent and subsequently migrated westward. It appears that only during the Greek period did sesame became a major oil-producing crop in south-west Asia.

C H A PTER 6

Fruit trees and nuts

Fruit-bearing trees constitute an important element of food production in the countries bordering the Mediterranean Sea. Their long-standing economic importance is amply reflected in classical traditions (Stager 1985). Five of the Biblical ‘seven species’ are founder crops of Mediterranean agriculture. Olive oil, wine, raisins, dates, and common figs were (and still are) staple agricultural products in south-west Asia and the Mediterranean basin. Like the Neolithic grain crops, but significantly later, the first indigenous fruit trees were brought into domestication in the Fertile Crescent area (Zohary and Spiegel-Roy 1975). In comparison to grain agriculture, horticulture evolved much later in the history of food production in Europe and southwest Asia. The first definite signs of fruit tree domesticants appear in south-west Asia in Chalcolithic contexts (seventh millennium BP), that is, several millennia after the firm establishment of grain agriculture in this region. In Early Bronze Age, olives, grapes, and figs emerge as important additions to cereals and pulses throughout the eastern Mediterranean basin, including the Aegean belt, while date palms have been grown at that time in the warmer and drier southern niches of this region. Horticulture is very different from grain crop agriculture. Cereals and pulses are ‘short investment’ annual crops. Their mature grains are ready to be harvested several months after sowing. The grain grower can move from place to place after the harvest, and practice shifting farming. In contrast, fruit trees are perennials. Orchards start to bear fruit three to eight years after planting, and reach full productivity several years later. Horticulture, therefore, needs protection from intruders the year round, indicating a settled way of life. 114

Genetically, domestication of fruit trees means changing the reproductive biology of the plants involved (Zohary and Spiegel-Roy 1975; Zohary 1984) by shifting from sexual reproduction (in the wild) to vegetative propagation (under cultivation). As a rule, domesticated varieties of fruit trees are maintained vegetatively by the farmer (as clones) by rooting of twigs, use of suckers, or by the more sophisticated technique of scion grafting. They are seldom raised from seed. This is in sharp contrast with their wild relatives, which reproduce mainly from seed. In other words, wild fruit tree populations maintain themselves through sexual reproduction and are distinctly allogamous. Crosspollination is brought about either by self-incompatibility or by dioecy (separate male and female individuals). Spontaneous populations manifest wide variability and maintain high levels of heterozygosity. Consequently, their seedlings segregate widely in numerous traits, including the size, shape, and palatability of the fruits. In the hands of the farmer, vegetative propagation has been a powerful device to prevent genetic segregation and to ‘fix’ desired types. By discarding sexual reproduction and inventing clonal propagation, the farmer can: (i) select and maintain exceptional individuals with desirable fruit traits from among large numbers of variable inferior trees, and (ii) duplicate (clone) the chosen types to obtain genetically identical saplings. In the case of fruit trees, this is no small achievement. Since fruit trees are generally cross-pollinated and widely heterozygous, most progeny obtained from seed (even progeny derived from the superior cultivars) do not bear the desired traits and therefore are economically worthless. The shift from seed planting to

FRUIT TREES AND NUTS

vegetative propagation has been the practical solution to assure a dependable supply of desired genotypes. In most fruit trees, this development made domestication possible. Only several nut trees (such as almond, walnut, and carob) were traditionally maintained by seed planting. Plant remains retrieved from archaeological excavations indicate that the olive, grapevine, fig, date palm, pomegranate, and the sycamore fig were the first fruit trees introduced into domestication in south-west Asia and Europe. Significantly, all these ‘first-wave’ fruits lend themselves to simple vegetative propagation by cutting and rooting of twigs (in the grapevine, fig, and sycamore fig), digging out of suckers (in pomegranate), planting basal knobs (in olive) or by transplanting offshoots (in date palm). In all these founder fruit crops the early growers did not have to resort to more sophisticated techniques of vegetative propagation (such as grafting). Wild olives, grapes, figs, dates, and pomegranates were thus ‘pre-adapted’ for early domestication. This adaptation was probably vital for their success in becoming the first domesticated fruit trees in south-west Asia and the Mediterranean basin. Several other fruit trees, such as apple, European pear, plum, sweet cherry, carob, and pistachio were introduced into cultivation much later. Definite evidence for their domestication appears only in the third millennium BP and their extensive incorporation into horticulture seems to have taken place only in Greek and Roman times. A plausible explanation for the late appearance of these ‘secondwave’ domesticated fruits is that they do not lend themselves to simple vegetative propagation. Their maintenance is based almost entirely on grafting. The Greeks and the Romans were already familiar with this art, and we have ample documentation that in classical times, apples and pears were maintained by this sophisticated method of vegetative reproduction (for review, see White 1970, pp. 248, 257–8). It is not clear when and where detached scion-grafting was invented. The earliest reliable description of grafting in the Mediterranean basin was given by Theophrastus (II, v. 3) who lived in Greece in the fourth century BC. Obviously, the introduction of this sophisticated method of propagation enabled the domestication of a new range of fruit trees. Grafting was probably invented outside

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the region of Mediterranean horticulture, and introduced into this region from the east. The earliest information on the use of grafting comes from China in connection with citrus-fruit domestication, and one of these texts is dated to ca. 139 BC (Cooper and Chapot 1977; Métailié (2007). The adoption of clonal cultivation means that most fruit trees have undergone very few sexual cycles in the five or six millennia since their introduction into cultivation. Domesticated clones persisted for hundreds of years. From the standpoint of evolution under domestication, this means a severe restriction on selection. In other words, selection (both conscious and unconscious) could have operated only during a limited number of generations, and we have to expect that the cultivars have not diverged considerably from their progenitors’ gene pools. This is in sharp contrast to the annual selfpollinated grain crops, where selection could have operated effectively during thousands of generations. Indeed, the domesticated varieties of fruit trees can be regarded as exceptional, highly heterozygous individuals of their biological species— clones that excel primarily in fruit size and fruit quality. The absence of profound genetic changes in the fruit trees under domestication is also apparent in their ecology. The climatic requirements of the cultivars closely resemble those of their wild relatives. Unlike the cereals and pulses, the fruit crops have not been pushed much beyond the climatic requirements of their wild ancestors. As already stated, most fruit trees under domestication are derived from wild progenitors in which cross-pollination is maintained by one of the following genetic systems: self-incompatibility, dioecy, or dichogamy. Because of this background, the shift from sexual reproduction to planting of vegetatively propagated clones introduced serious limitations on fruiting. Planting of a single self-incompatible clone, or alternatively female clones, would not bring about fruit set. Several agronomic devices, assuring fruit set in the orchard, have been empirically adopted. They were accompanied by unconscious selection for several types of mutations that resolved the restrictions set by self-incompatibility and sex determination. In hermaphroditic, self-incompatible species such as olive, apple, or pear, the early planters very likely

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realized that to obtain satisfactory fruit set, it was necessary to plant together two or more synchronously-flowering clones (Table 8). The traditional cultivation of such fruit trees is based on mixed planting, a practice which brings about pollination between different genotypes. Two additional solutions are the outcome of unconscious selection: (i) in the peach, apricot, sour cherry, European plum, as well as in several cultivars of almonds or olives, we find mutations that caused the breakdown of selfincompatibility, (or at least rendered the system ‘leaky’), so that self-pollination also results in considerable fruit set in such clones; and (ii) in other self-incompatible taxa, pollination has been dispensed with altogether by incorporation of mutations conferring parthenocarpy (fruit development without fertilization and without seed set). Several clones of cultivated pear show this adaptation. In dioecious species, fruit set is safeguarded in parallel ways. In the pistachio, ‘mixed planting’ is employed, and some male individuals are planted together with the female clones. In date palms and Smyrna-type figs, natural pollination is frequently augmented by artificial pollination. Also in dioecious fruit trees, the early planters must have unconsciously selected two types of mutations that rendered cross-pollination unnecessary: (i) a genetic Table 8 Horticultural safeguarding fruit set in fruit trees (compiled after Zohary 1984) The Genetic System The Agricultural Solution Self-incompatible fruit trees

1. Planting together a few synchronously flowering clones. 2. Mutations toward breakdown (some almonds, 4x cherries) or reduction (e.g. some olives, plume, apples, and pears) of self-incompatibility. 3. Parthenocarpic-induced mutations (e.g. some pears).

Dioecious fruit trees

1. Planting additional male plants (e.g. pistachio). 2. Artificial pollination (e.g. date palm, Smyrna-type figs). 3. Unconscious selection toward: (i) hermaphrodite plants (e.g. grape) (ii) parthenocarpy (e.g. common fig, Corinth-type grapes).

change from dioecy (in the wild) to hermaphroditism (under domestication) evolved in the grapevine; and (ii) a replacement of sexual reproduction by parthenocarpy occurred in many clones of the fig and the sycamore fig. In most fruit trees, population variation in seed morphology is quite large. This phenomenon makes definite differentiation between wild and domesticated forms in early archaeological contexts problematic. Clear-cut signs, equal to the smooth/rough disarticulation scar in the cereals (p. 22) are currently unknown. Like cereals, such differences, especially in seed-size, can be seen in later phases. As a result, we cannot be certain whether the earliest finds are wild or domesticated. Recent adoption of morphometric tools (such as Terral et al. 2010) might lead to better identification in the future.

Olive: Olea europaea The olive, Olea europaea L. (Oleaceae), is the most prominent, and economically the most important classical fruit of the Mediterranean basin (Zohary 1995a). It comprises the oldest group of plants that founded horticulture in the Old World, together with grapevine, fig, and date palm (Zohary and Spiegel-Roy 1975; Boardman 1976; Stager 1985). Since Bronze Age, the wealth of many Mediterranean peoples centred on the cultivation of olives, which provided valuable storable oil as well as edible fruits. Olive oil has been used in eating and cooking, as well as for ointment and lighting. Because of its excellent durability, it served as a principal article of commerce. The whole fruit was also preserved and consumed. Bread and olives were, and still are, a staple diet in peasant communities throughout the Mediterranean basin. Olives grow in typical Mediterranean climates. This fruit crop, and its closely related wild oleaster forms, are considered reliable indicators of Mediterranean environments. The olive is a relatively slow-growing tree, diploid (2n = 4x = 46 chromosomes), and fruit production starts five to six years after planting. If well managed, the longliving olive trees can keep fruiting for hundreds of years. Olives, under domestication, manifest considerable variation in the size, shape, and oil content of their fruits. Hundreds of distinct varieties

FRUIT TREES AND NUTS

are recognized. Different parts of the Mediterranean basin are frequently characterized by specific local forms. The olive crops can be roughly subdivided into two main types: (a) oil varieties, the ripe fruits of which contain at least 2% oil (the oil is monounsaturated with a high percentage of Oleic acid and used for cooking, salt dressing, and food preservation); and (b) table olives: less oily forms used for the preservation of whole fruits, by pickling or salting. There are also dual-purpose cultivars (see Lavee and Zohary 2011). Carbonized kernels (stones) and charred wood of pruned twigs, used as firewood, constitute the bulk of olive remains in archaeological excavations.

Wild ancestry The domesticated olive, O. europaea L., shows close affinities to a group of wild and feral olives distributed over the Mediterranean basin and traditionally referred to as ‘oleaster’ olives (Plate 13). These wild forms are fully inter-fertile with the domesticated varieties, and interconnected with them by sporadic, spontaneous hybridization (Zohary and Spiegel-Roy 1975; Zohary 1995b; Lavee and Zohary 2011). Oleaster olives and the domesticated clones have similar climatic and soil requirements. Previously, many botanists regarded the wild oleaster forms as representing an independent species, O. oleaster Hoffm. & Link. More recently, because of its close morphological and genetic affinities to the domesticated fruit tree, most researchers dealing with Mediterranean plants regard the oleaster forms as the wild progenitor of the crop. They place it within O. europaea L. species complex, either as a subspecies [subsp. oleaster (Hoffm. & Link) Hegi] or as a variety [var. sylvestris (Mill.) Lehr. = var. oleaster (Hoffm. & Link) DC]. Oleaster olives differ from the domesticated clones mainly by their smaller fruits and usually by spinescent juvenile branches. These wild fruits have less fleshy mesocarp and contain less oil. However, frequently the stones are not much smaller than those in the cultivars. Wild olives are almost fully selfincompatible, and as with many other trees maintaining such a genetic system, oleaster populations often show a wide range of electrophoretically discernible allozyme variation. In comparison, the

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domesticated varieties (of the domesticated gene pool) are much less variable (Quazzini et al. 1993; Zech-Matterne and Leconte 2010). Under cultivation, the reproductive biology of the tree changed. While wild olives reproduce by sexual reproduction from seed, domestic varieties are maintained by vegetative propagation and are, in fact, clones. Traditional vegetative propagation depends primarily on the utilization of knobs (= uovuli) and truncheons that develop at the base of the trunk, and by grafting. Today, vegetative propagation in horticulture relies mainly on tissue culture. In Olea (as with numerous other fruit trees) the shift to vegetative propagation is the cultivator’s countermeasure to the problems of wide genetic segregation that characterize reproduction in crosspollinated plants. Domesticated olive clones are considerably heterozygous. When their sexually produced seeds are planted, the ensuing progeny segregate widely. In fact, most sexual seedlings resemble the wild forms in their morphology and are poor or useless in terms of fruit quality. Consequently, propagation from seed is impractical in oleoculture. In order to ‘fix’ useful genotypes, the grower has to resort to clonal propagation. Seedlings can only be used as a variable raw material for selection of new clones. At present, such selection of seedlings is performed in some olive-breeding programs. Spontaneous seedlings have accompanied olive growing from its very start, and rare individuals showing superior qualities may have caught the attention of early cultivators and were then picked up as the new clones. Over large areas in the Mediterranean basin, wild oleaster-type, olives thrive as common constituents of maquis and garrigue formations (Map 15). In addition, these shrubs and small trees frequently colonize secondary habitats, such as the edges of cultivation or abandoned orchards. In such places, particularly in areas of olive plantations, spontaneous olives seem frequently to be ‘feral’. Such ‘feral’ plants can derive from hybridization (i) between different domestic clones, or (ii) between domesticated clones and adjacent wild oleaster plants. The Mediterranean wild olives are associated with olive cultivation also in another way. They often served as stock material for grafting. Oleaster suckers or knobs are being dug out (in the wild),

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Fig. 34 Charred stones of olives, Olea europaea, Chalcolithic Tuleilat Ghassul, Jordan (Zohary and Spiegel-Roy 1975).

and planted in orchards as hardy stock material for grafting of domestic scions. In some localities, for example in western Turkey, one still encounters another old tradition, namely the grafting of wild olives growing in non-arable maquis and/or garrigue vegetation. The grafted wild olives in such niches are protected by the farmers, while other trees and shrubs are cut for firewood and further suppressed by intensive grazing. Olea europaea is the only Mediterranean representative of the genus Olea L., which includes some

two to twenty-five species distributed over tropical and southern Africa (the main center), and southern Asia and China (as well as eastern Australia, New Caledonia, and New Zealand). Both the olive crop and the Mediterranean oleaster forms are genetically closely related to (and probably inter-fertile with) several non-Mediterranean wild olive taxa. Most widespread among the latter are various east African, south Arabian, south Iranian, and Afghan wild subspecies of the O. europea taxonomic complex (subsp. sylvestris, subsp. cerasiformis, subsp.

FRUIT TREES AND NUTS

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0 100 200 miles 0

200

400 km

Map 15 Geographical distribution of wild olive, Olea europaea subsp. oleaster (based on Zohary and Spiegel-Roy 1975; Lavee and Zohary 2011). Additional wild forms of the olive species-complex occur in: (i) southern and east Africa (including west and south Arabian Peninsula); (ii) Afghanistan; and (iii) west China (Zohary 1994; Lavee and Zohary 2011). Note that only the wild Mediterranean oleaster stock seems to be the progenitor of the domesticated olive.

laperrinei, subsp. cuspidata, subsp. ferruginea, subsp. chrysophylla, subsp. africana). The morphological differences between these more tropical wild olives and their Mediterranean counterparts are relatively small. Consequently, Green and Wickens (1989) regarded some of them only as an additional subspecies of the European olive and named them O. europaea L. subsp. cuspidata (Wall.) Ciferri. However, these south Asian and east African olives are geographically separated from their Mediterranean relatives by wide morphological and molecular discontinuities. They are also adapted to different environments. Such isolation may justify their ranking as independent species. Using multiple types of molecular markers (e.g. mitotypes, SSR, and RAPD data), Breton et al. (2003, 2006) concluded that eastern and western Mediterranean oleaster populations differ, with cultivars of eastern origin (populations from Israel, Syria, and Turkey). Also, they suggest at least two simultaneous domestications occurred on opposite ends of the Mediterranean Basin, possibly the second one in the Spain/Corsica region. Other non-Mediterranean wild types closely related to the crop are O. europaea subsp. laperrinei

(Batt. & Trab.) Cif. and O. europaea L. subsp. cerasiformis (Webb & Berth.) Kunkel & Sunding [= O. europaea L. var. maderiensis Hart.]. The first is a Saharo-Montane relic that bridges the Mediterranean forms with their African savannah relatives. It is confined to a few inner mountains in the Sahara Desert and does not come close to the Mediterranean forms, except perhaps in the southern Atlas Mountains. The second is the wild olive of the Macronesian Islands. (For details on the taxonomy and distribution of all these non-Mediterranean wild olives consult Green and Wickens 1989; Zohary 1994; Lavee and Zohary 2011). The evidence from living plants therefore shows that the domesticated olive is closely related to the Mediterranean oleaster wild forms. These could be regarded as the general stock from which the domesticated fruit tree had been derived.

Archaeological evidence It is likely that olives were collected from the wild long before their domestication. The olive find in Epi-Palaeolithic Ohalo II, Sea of Galilee (Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009; Weiss

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et al. 2004, 2008) undoubtedly represent gathering from the wild. Some olive stones were found in ca. 8,150–7,850 cal BP Pottery Neolithic Nahal Zehora, Mount Carmel, Israel (Kislev and Hartmann forthcoming). Also the thousands of waterlogged stones (and stone fragments) uncovered in the submerged, late Neolithic (ca. seventh millennium BP) sites off Mt Carmel (Galili et al. 1989; Galili and Shavit 1994– 5) seem to represent gathering from the wild as well (see Kislev 1994–5). The numerous charred stones found in contemporary Dhali Agridhi, Cyprus (Stewart 1974), are very likely of a similar nature. So are the stones and/or charcoal recovered from the Mesolithic and Neolithic Grotta dell’ Uzzo, Sicily (Costantini 1989) or from Neolithic and Chalcolithic sites in Spain (Buxó 1997), and in Natufian and early Neolithic Nahal Oren, Mt Carmel (Noy et al. 1973). Definite signs of olive domestication come from several Chalcolithic sites in Israel and Jordan. Numerous well-preserved, carbonized olive stones (see Fig. 34), as well as charred olive wood, were uncovered in ca. 6,800–5,800 cal BP Chalcolithic Tuleilat Ghassul, north of the Dead Sea, together with cereal grains, dates, and pulses (Zohary and Spiegel-Roy 1975; Kislev 1987; Neef 1990). This site lies far outside the natural range of wild olives. No oleaster olives occur today in the lower Jordan Valley, or below the sea-level escarpments. The region is too dry for it, which was probably the case in Chalcolithic times. In Israel and Jordan, the areas nearest to the lower Jordan Valley that support wild olives are the western flanks of the Judean Hills, Mt Carmel, and the Gilead. This indicates that the Tuleilat Ghassul olives may have been products of cultivation. They were probably raised under irrigation in a similar manner in which olives are grown today in the Jordan Valley. The finds of rich olive remains in Tuleilat Ghassul are supplemented by olive stones and charcoal remains from: (i) three additional Chalcolithic sites in the Jordan Valley, namely Tell Saf (Gophna and Kislev 1979), Tell Shuna North, and Tell Abu Hamid (Neef 1990)—they contain masses of fragments of crushed stones; i.e. waste products of olive pressing; and (ii) contemporary sites in the Golan Heights (Epstein 1978, 1993). Additional support is given by a simultaneous marked rise in olive pollen values around 6,500 cal BP in the pollen dia-

gram of Birkat-Ram, Golan Heights (Neumann et al. 2007). Later, olive remains abound in ca. 4,900–4,700 cal BP Early Bronze Age Arad (Hopf 1978c), Bab edhDrah (McCreery 1979), ca. 4,900 cal BP Tell esSa’idiyeh, Jordan (Cartwright 2003; and pers. comm.), and several other Early Bronze Age sites (Liphschitz et al. 1991; Liphschitz 2008). Outside Israel and Jordan, Chalcolithic and Early Bronze Age finds of olives have been few thus far. Olive remains were found in fifth millennium BP Tell Soukas, Syria (Helbaek 1962), in ca. 4,850–4,200 cal BP Troy, Turkey (Riehl 1997), and in Early and Middle Bronze Age Marki Alonia (Adams and Simmons 1996), and ca. 3,450–3,350 cal BP Hala Sultan Tekke (Hjelmqvist 1979a), Cyprus. A few are also present in several early Minoan sites such as Myrtos (Renfrew 1972), and Knossos, Crete (Evans 1928, p. 135). Some stones were found even from ca. 5,400–4,200 cal BP Zambujal, Portugal (Hopf 1981), where other sources of evidence (Blitzer 1993) indicate olive cultivation at this time. Sarpaki (Sarpaki 2009) indicates, however, that olives are totally missing from Knossos. In the Middle and Late Bronze Age, olive cultivation and olive-oil production seem already to have been well established throughout the countries bordering the east shore of the Mediterranean Sea (Stager 1985) and in Late Bronze Age, also in mainland Greece (Boardman 1976; Hansen 1988). The successful establishment and large scale utilization of Olea is indicated also by the increase, from early Bronze Age on, of olive pollen grains in cores obtained from the Sea of Galilee (Baruch 1990), as well as by the appearance of numerous presses, olive oil vessels, and depiction of olives in Bronze Age art. Apparently, olive horticulture did not play a major role in Bronze Age Egypt. Instead, the export of olive oil from the southern Levant to Egypt is well documented in the Bronze Age archaeology (Stager 1985). Although charred olive stones were found already in ca. eighteenth to the sixteenth centuries BC Kom el-Rabi’a, Memphis (Murray 2000b) and Tell el-Dab’a, Nile Delta (Thanheiser 2004), as cited by (Newton et al. 2006), it is hard to determine their origin. They were most probably imported. Likewise, the unique finds of olive leaves and twigs in garlands and jars of olive oil from ca. 1,325 BC

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Tutankhamun tomb (Germer 1989b; Hepper 1990; de Vartavan et al. 2010). Olive cultivation was probably introduced into the west Mediterranean basin in the early part of the third millennium BP by the Phoenician and Greek colonists (Boardman 1976). This trade is well demonstrated by thousands of olive stones, including some 2,500 in a single Canaanite Jar, from late fourteenth century BC Late Bronze Age Uluburun shipwreck off the southwestern Turkish coast (Haldane 1993). In conclusion, in the Mediterranean basin olives constitute a complex of wild forms, weedy types, and domesticated clones. All are genetically loosely interconnected and show similar climatic and edaphic preferences. Information from the living plants corresponds well with the available archaeological evidence. The earliest archaeological records of olive cultivation come from the lands bordering the eastern shores of the Mediterranean Sea—a territory where genuinely wild oleaster olives thrive today. Hence, on the basis of the combined evidence, one is led to the conclusion that the olive was probably first brought into cultivation in the Levant.

Grapevine: Vitis vinifera Grapevine, Vitis vinifera L. (Vitaceae), is one of the classical fruits of the Old World. Together with the olive, common fig, pomegranate, and date palm, it comprises the oldest group of fruit trees around which horticulture evolved in the Mediterranean basin (Zohary and Spiegel-Roy 1975). Since Early Bronze Age, grapes have contributed significantly to food production in this area, providing fresh fruits rich in sugar (the berries contain 15–25% sugar), easily storable dried raisins, and juice for fermentation of wine. The latter became an important trade element in the countries around the Mediterranean Sea. The grapevine thrives in Mediterranean-type environments, but, compared to the olive, it tolerates cooler and more humid conditions. For this reason, viticulture extends northward beyond the Mediterranean basin and succeeds in areas with a relatively mild climate in western and central Europe, and in western and central Asia. Vitis is a perennial, diploid (2n = 2x = 38 chromosomes)

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climber. Wild populations of Vitis are dioecious and reproduce from seeds. Under domestication, grapevine cultivars are propagated vegetatively by rooting winter dormant twigs or by grafting. The grapevine has to be pruned yearly in order to remain confined to a manageable size, and for regulation of fruit production. As with most other fruit trees, viticulture is based on the ‘fixation’ and maintenance of vegetative clones. Almost all grape cultivars bear hermaphrodite flowers and set fruit by self-pollination. Traditional Old World viticulture was based on thousands of distinct clones (Alleweldt et al. 1991; Mullins et al. 1992; Olmo 1995b). These vary widely in their habit, climate, and soil preferences, as well as in the shape, size, colour, taste, and sweetness of their fruits. Juicy, small-berried varieties with a rather acid taste are commonly used for wine production, especially in Europe. Types with sweet, large fruits prevail as table grapes. Some clones, such as the traditional Black Corinth and Sultanina, bear small, seedless berries appreciated in raisin production. The berries of many of tablebred varieties are seedless. The grapevine is a fast growing fruit crop. Production usually starts three years after planting. Charred pips and wood constitute the bulk of grape wine remains in archaeological excavation, although some sites also yield whole fruits as well.

Wild ancestry The genus Vitis contains some sixty-five species native to northern and south-east Asia and central America. All species are diploid, and are largely self-pollinated. The domesticated grapevine, V. vinifera L., is closely related to an aggregate of wild and feral forms (Fig. 35a) distributed over Europe and western Asia. Classical botanists regarded these wild forms as an independent species, V. sylvestris (C.C. Gmelin) Hegi. However, since these wild forms show close morphological and genetic affinities with the domesticated stock, most botanists now regard sylvestris as the wild race of the domesticated crop. Currently, they are classified as subspecies sylvestris (C.C. Gmelin) Berger, together with subspecies vinifera, within the V. vinifera crop complex, and are considered as the source from

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which the cultivars could have been derived (see, for example, Hegi 1935; Webb 1968; Mullins et al. 1992). These wild forms are largely dioceous, diploid, and their population contains 50% male and 50% female individuals. In such wild populations, cross-pollination is the rule. Sylvestris grapes are widely distributed from the Atlantic coast to Tadzhikistan and Kazakhstan (Map 16). A unique community of wild grapes grows in southern Kazakhstanthe most northern area of wild grapes in central Asia (Dzhangaliev et al. 2003). They are primarily forest climbers, suited to the humid and climatically mild deciduous forest area south of the Caspian Sea and along the southern coast of the Black Sea. Sylvestris grapevines also abound in the relatively cooler and more mesic northern parts of the Mediterranean sclerophyllous vegetation belt—from Turkey and Crimea, through Greece and former Yugoslavia, to Italy, France, Spain, and north-west Africa. Along the Rhine and the Danube, wild grapes used to penetrate deeply into central Europe, yet these populations are largely extinct. Scattered stands of wild grapes occur also in more xeric territories and in less woody places in the Fertile Crescent. However here, sylvestris grapes are mostly confined to gorges and to the vicinity of springs and streams. The boundary between the domesticated grape clones and the wild forms is blurred by the presence of escapees and secondary derivatives of hybridization. Spontaneous crossing between wild plants and cultivars have been found repeatedly where sylvestris grapes grow in close proximity to vineyards, as the F1 hybrids are fully fertile. To summarize, in V. vinifera we are faced, (in the Mediterranean basin and in south-east Asia) with a variable complex of wild forms (growing in primary habitats), escapees and seed-propagated weedy types (that occur mainly in disturbed surroundings), and orchards of cultivars. The picture of the pre-agriculture distribution of the wild grapevine has probably been blurred by introgression between tame and wild types, and by ‘weedy’ forms occupying human disturbed environments. However, there can be little doubt that sylvestris grapes are indigenous elements in southern Europe, the Euxinian and the Hyrcanian vegetation belts south of the Black Sea and the Caspian Sea, and relatively wet places in

the Mediterranean vegetation belt in south-west Asia. Sylvestris grapes differ from the domesticated varieties by their relatively small and usually acid berries (Fig. 35a), which are quite suitable for the preparation of wine (for details see Olmo 1995a, Table 3.1). Wild grapes may also be recognized by more numerous (three to four) pips per berry, and by somewhat more globular pips, generally with stalks or ‘beaks’ constricted at the attachment to the main body of the pip (Fig. 35b). Among the thousands of domesticated grapevine clones and among the variable wild sylvestris populations, the range of variation in pip morphology is very wide. Furthermore, the variation in the domesticated assemblage and in the wild forms overlaps considerably. For this reason, pip morphology cannot be regarded as a completely safe diagnostic trait for distinguishing between wild and domesticated Vitis remains in archaeological excavations (but see different view in Kislev 1988). Kroll (1999b), however, points to a difference in pip production between wild and domestic grapevines. As wild Vitis are usually dioecious, an unpollinated female flower will fail to produce seeds and will drop. In contrast (see below) nearly all

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Fig. 35a Fruiting wild grapevine, Vitis vinifera subsp. sylvestris (Zaprjagaeva 1964, plate 283).

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Map 16 Geographical distribution of wild grapevine, Vitis vinifera subsp. sylvestris (based on Zohary and Spiegel-Roy 1975). Wild grape extends toward the east beyond the boundaries of this map and reappears in a few locations in Turkmenistan, Tadzhikistan, and Kazakhstan.

contain three to four fully developed pips, but never underdeveloped pips. On the other hand, the berries of (traditional) domestic varieties frequently contain two normal sized pips and a partly developed third pip. The presence of such reduced pips may indicate domestication.

domesticated grapes are hermaphrodites, and selfpollination in their flowers is possible. Such a flower may develop in several directions. It can drop undeveloped, develop into a berry with underdeveloped pips, or develop into a berry with no pips (‘seedless’ varieties). For this reason, berries of wild forms (b)

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Fig. 35b Variation in pip morphology in wild forms of the grapevine, Vitis vinifera subsp. sylvestris (Zaprjagaeva 1964, Plate 286). Note the wide variation in the size and shape of the pips and their beaks.

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Domestication has brought about considerable changes in the reproductive biology of the grape. The first obvious change is the shift from the wildtype sexual reproduction (by seed) to vegetative propagation of clones (by cuttings or by grafting). Just as in the olive, this is the cultivator’s way of overcoming wide segregation in the seedlings and achieving ‘fixation’ of desired types. The second conspicuous development is the breakdown of the wild-type system of sex determination. Usually, wild sylvestris plants are dioecious, and their populations contain an equal proportion of male and female individuals. Fruit setting in the wild depends on cross-pollination. In contrast, nearly all domesticated grape varieties are hermaphrodites (self-fertile). Their flowers contain both a pistil and anthers. The hermaphroditic condition ensures self-pollination and fruit setting without the need of male pollen donors. Sex determination in wild sylvestris populations is governed by a single gene (Olmo 1995b). Female individuals are homogametic—they carry a homozygotic recessive genotype Sum Sum that suppresses the development of anthers. Male individuals are heterozygous for a dominant, female suppressing SuF allele, and have a SuF Sum genotype. The change, under domestication, from bisexuality (= dioecism) to hermaphroditism was attained by a mutation shift to allele Su+ that is dominant over Sum and brings about the development of both the pistil and the anthers. Many hermaphroditic cultivars are still heterozygous and posses a Su+ Sum genotype. Other hermaphroditic clones contain a homozygous Su+ Su+ genotype. Vitis vinifera is the sole Mediterranean representative of the genus Vitis. This is a rather large and variable genus comprising some sixty species (de Lattin 1939; Levadaux 1956; Mullins et al. 1992). About two-thirds of these species occur in temperate North America, and one-third is distributed over temperate and subtropical parts of East Asia. All wild members of the genus Vitis are dioecious, perennial woody climbers with coiled tendrils. All known Vitis species, (excluding the Muscadinia group which is now regarded as an independent genus) have 2n = 2x = 38 chromosomes, and can be easily crossed experimentally. Their F1 hybrids are vigorous and fertile. In nature, the principal taxa (‘species’) are often not fully reproductively isolated

from one another by various geographical and ecological barriers. Consequently, the boundaries between the main morphological forms are frequently blurred, making a clear-cut delimitation of species in Vitis a difficult or even an impossible task. Before the colonization of North America, viticulture in Europe and western Asia was restricted to the gene pool present in the V. vinifera complex. However, in modern times, several wild species native to America have been used either as additional genetic sources for breeding of new varieties of grapes, or as hardy wild sources of stocks conferring resistance against the devastating attack of the Phylloxera root aphid.

Archaeological evidence Berries of wild Vitis were gathered from the wild long before the domestication of the V. vinifera. Charred pips and occasionally also entire berries or raisins have been discovered in numerous prehistoric sites in Europe and south-west Asia, particularly in northern Greece, former Yugoslavia, Italy, Switzerland, Germany, France, and Spain (for enumeration of finds, see Riviera Núñez and Walker 1989; Kroll 1991; Colledge and Conolly, 2007). Several dozens pips discovered in late Neolithic (ca. 9,000–7,600 cal BP) Dhali Agridhi, Cyprus (Stewart 1974) also seem to represent wild grape. In some of these prehistoric sites, like ca. 8,200–6,650 cal BP Anza, Macedonia (Renfrew 1976), ca. 7,800–7,600 cal BP Knossos, Crete (Sarpaki 2009), and ca. 6,350– 6,150 cal BP Tell Hârşova, Rumania (Cârciumaru 1996; Monah 2002; Monah and Monah 2008), such pips were found together with domesticated grain crops. As already mentioned (see p. 122) the range of variation in pip morphology in domesticated and in wild forms overlaps considerably. For this reason pip morphology cannot be regarded as a fully safe diagnostic trait for distinguishing between wild and domesticated Vitis remains in archaeological excavations (and see different view in Kislev 1988). However, the appearance of a crop in an ecological environment where its wild relative does not grow can support the argument for its domestication

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status (Zohary and Spiegel-Roy 1975, and recent review in Miller 2008). The earliest probable domesticated finds of V. vinfera come from several Early Bronze Age sites in the Levant. Pips and charred berries containing two to three seeds were uncovered in Early Bronze Age (ca. 3,500–3,300 cal BP) Jericho, Israel (Hopf, 1983). These finds have been complemented by remains found in Early Bronze Age Lachish (Helbaek 1958) and Arad (Hopf 1978c), Israel, Numeira, Bab edhDhra (McCreery 1979), and ca. 4,900 cal BP Tell esSa’idiyeh (Cartwright 2003; and pers. comm.), Jordan, and Kurban Höyük, south-eastern Turkey (Miller 1986). Arad yielded two samples of charred wood, and in Numeira numerous pips and hundreds of whole berries were excavated. The fruits are small, and the pips roundish and relatively short-beaked. But since wild Vitis is absent today in the Jordan Valley and Judea, and is unlikely to have grown wild in these areas in the second half of the sixth and the fifth millennia BP, the combination of pips and charred wood provides a solid proof that Vitis was already domesticated at that time. Additional archaeological finds are available from sites in south-east Turkey and in North Syria (for enumeration of sites see Zettler and Miller 1995; Miller 2008). The remains retrieved from Kurban Höyük, Urfa district, south Turkey (Miller 1986; Miller 1991, 2008) are instructive since pips of grapes were recorded from 5% of the samples taken from late Chalcolithic layers in this site. Their occurrence increased to 10% of the samples taken from Early Bronze Age levels. They became frequent (66%) and appear in masses in mid-late Early Bronze Age. As reported by Miller (1982), mineralized pips, presumably from cess deposits, are the most direct evidence of fruit consumption in late fifth millennium latrines at the site. This profile seems to reflect a rapid growth of viticulture in the north Levant at the start of the Bronze Age. If confirmed, the charred pips from Chalcolithic Tell Shuna North in the Jordan Valley (Neef 1998), might represent an even earlier domestication. Finally, it was suggested that infrared spectroscopy detection of tartaric acid in residues on archaeological vessels is a marker for wine (Singleton 1994). As Miller (2008) notes, the chemical signature of tartaric acid can result from various other related

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products: vinegar, raisins, grape juice, pekmez (grape ‘molasses’), or one of few other plants that produce it (see Cavalieri et al. 2003). At present, the earliest evidence of tartaric acid comes from the Neolithic site, ca. eighth millennium BP, Hajji Firuz, Turkey (McGovern et al. 1996; McGovern 2003). As this site lies at the edge of the current range for wild grape (Map 16), it is hard to determine whether these were wild or domesticated grapes. Such residues were found on the inner surface of a large jar recovered from sixth millennium BP Godin Tepe, western Iran, some 400 km distant from wild grapevine territory (Badler 1995; Badler et al. 1990; McGovern and Michel 1995). From the second half of the fifth millennium BP on, fresh grapes, raisins, and wine are also repeatedly recorded in Mesopotamian cuneiform sources (Postgate 1987). Grape growing areas in the south Levant were exporting wine and raisins to Egypt already in the Early Bronze Age (Stager 1985; James 1995). The dimensions of this trade were demonstrated further recently by a discovery of hundreds of vessels at the tomb of one of Egypt’s earliest kings, buried around 5,100 BP at Abydos (McGovern 1998). Neutron activation analysis of jars showed that they were made in south Levant. The infrared spectrometry tests of sediments left inside the jars identified tartaric acid, and confirmed the suspicion that they were wine vessels. This conclusion was further supported by a few pips found inside some of the jars. Somewhat later, during the first and second dynasties (ca. 5,000–4,700 BP), signs appear that viticulture and wine making were introduced into the Nile Valley. Remains of grapes, depiction of grapevine growing, wine production, and evidence on import of wine and raisins continue to appear in Egypt from the Old Kingdom times onwards (Germer 1989b; James 1995; Murray 2000c). Among the earliest finds are from Pre-dynastic Tel Ibrahim Awad (de Roller 1992; Thanheiser 1992) and Tell el-Fara’in (Buto), the Nile Delta (Thanheiser 1991), and a rich find of fragments of desiccated raisins in third dynasty, sometime around 4,600 cal BP, Djoser pyramid at Saqqara (Lauer et al. 1950; Germer 1985). To the east of Egypt, the earliest evidence currently comes from ca. 2,200–1,600 cal BP Jerma (Garama), Libya (Pelling 2008).

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Viticulture does not flourish in most parts of the hot Nile Valley. As argued by Stager (1985), the most plausible source for such introduction is the Levant. Climatically, Egypt lies outside the optimal range for Vitis. In dynastic times, raisins and wine were largely imported into Egypt, grapes were grown only as a luxury crop, and were restricted mainly to the cooler delta area. In the Aegean area signs of grapevine cultivation appear somewhat later. In Thessaly and Macedonia pips become so common at several late Neolithic (late second half of seventh Millennium onward BP) sites that Kroll (1991) remarks that their sheer numbers could suggest cultivation. However, convincing evidence comes only from Early Helladic IV (fifth millennium BP) Lerna in south Greece, where numerous pips were uncovered (Hopf 1961b). Additional remains were uncovered from Middle and from Late Bronze Age sites (Hansen 1988). They include several hundreds of pips from Late Helladic, second half of fourth millennium BP, Kastanas (Kroll 1983). Parallel to the situation in the Levant, the development of grapevine cultivation in the Helladic and Minoan cultures is indicated also by the presence of presses, and the appearance of specific wine jars and wine cups. Archaeobotanical documentation of Bronze Age V. vinifera is also available from Transcaucasia (Wasylikowa et al. 1991). Pips appear in Georgia in Early Bronze Age. They become common in Middle and Late Bronze Age (second and first millennia BC). Also in Armenia, pips were frequently retrieved from the second millennium BC onwards, while the earliest central Asian finds came from ca. 5,000–3,700 cal BP Anau, Turkmenistan (Miller 1999; Miller 2003). From the end of the fifth millennium BP, there are convincing signs of grapevine cultivation in Baluchistan. Grape pips were uncovered among food offerings placed in graves at ca. 4,000 BP Shahri-Sokhta, south-east Iran (Costantini 1977). Charred wood of Vitis (cf. V. vinifera) was retrieved from the latest strata (Period VII, ca. 4,000 BP) in Mehrgarh, Pakistan (Thiebault 1989), and pips were found in contemporary, neighbouring Nausharo (Costantini and Costantini-Biasini 1986), and in Miri Qalat (Tengberg 1999). Similar to Egypt, these territories lie far away from the wild habitats of V. vinifera.

These finds (particularly the charcoal) indicate imported cultivation. Viticulture was apparently introduced to the west Mediterranean basin by Phoenician and Greek colonists (Stager 1985; Buxó 1997). The Romans brought this crop to temperate Europe (Loeschke 1933; König 1989). In summary, the situation in Vitis parallels that found in Olea. The earliest archaeological indications of viticulture come from areas close to the eastern shore of the Mediterranean Sea. In this territory (Map 16), we still find wild sylvestris-type populations that might have been used for developing the cultivated grape. While oleaster olives are confined to the typical Mediterranean warm and summer-dry climate, sylvestris vines thrive in more temperate and mesic conditions. These adaptations are paralleled under cultivation. On the basis of the available information (both from the living plants and from archaeobotanical remains) we assume that the Levant is the probable area in which Vitis vinifera domestication could have been initiated.

Fig: Ficus carica The Mediterranean fig, Ficus carica L. (Moraceae), is the third classical fruit crop associated with the beginning of horticulture in the Mediterranean basin and south-west Asia (Zohary and Spiegel-Roy 1975). This common fig (on the basis of available evidence) has been part of food production in this area since Early Bronze Age, providing fresh fruit in summer and storable, sugar-rich, dry figs all year round. It is a relatively fast-growing fruit crop. Production starts three to four years after planting. Early fig cultivation was centred on typical Mediterranean environments, in close association with the olive and grapevine. The fig tree is diploid (2n = 2x = 26 chromosomes), functionally dioecious, cross-pollinated, and spontaneous populations show half-male and half-female individuals (Kjellberg et al. 1987). They depend entirely on sexual reproduction from seed. Under domestication their propagation is vegetative. The grower maintains desired genotypes by rooting of winter-dormant twigs and occasionally by grafting. Since F. carica is bisexual, female fruit-bearing clones are vegetatively propagated. Such cultivars are

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3 mm

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Fig. 36 Common fig, Ficus carica. A–a twig with a flowering syconium (‘fruit’) on top; B–Mature syconium; C–Cross-section in a syconium; D–Staminate flower with four anthers; E–Modified short styled female flower; F–Long styled female flower (with kind permission of J. Galil).

considerably heterozygous and when progenytested, they manifest wide segregation. Most of the seedlings are economically useless. The majority of present-day cultivars, known as ‘common figs’, are parthenocarpic. They were selected for their ability to produce sweet, fleshy, large ‘fruits’ (syconia) without pollination and fertilization. A second group of highly appreciated female cultivars, the ‘Smyrna-type’ figs, still require pollination and fertilization for fruit development (Condit, 1947; Zohary 1995a; Flaishman et al. 2008). A single dominant mutation determines the shift (under domestication) to parthenocarpy. The reproductive biology of F. carica is complex (Storey 1976; Galil and Neeman 1977; Valdeyron and Lloyd 1979; Beck and Lord 1988) and based on: (i) a highly specialized inflorescence (the syconium, Fig. 36) that the growers refer to as the fig’s ‘fruit’;

(ii) the presence of two sex forms of which the female morph is known as the ‘true’ or common fig, and the male as caprifig; and (iii) an elaborate symbiosis between the plant and its pollinator, the fig wasp Blastophaga psenes. The syconium (unique to the genus Ficus), is a fleshy flowering branch transformed into a hollow receptacle that bears numerous minute flowers on its inner surface, and is open to the outside by a narrow orifice (ostiole). The ‘true’ fruits are small druplets or ‘seeds’ each developing in a female flower inside the syconium. Young syconia on the ‘true figs’ contain only pistillate, female, long-styled flowers. In contrast male ‘caprifigs’ produce spongy, non-palatable syconia containing both staminate male flowers (Fig. 36D) and modified short-styled female flowers (Fig. 36E). Female flowers nourish the Blastophaga larvae by

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turning into galls when eggs are laid into them. True figs produce their main crops of ‘fruits’ in late summer. Caprifigs usually bear three crops of syconia during the year: over-wintering mamme, numerous profichi that develop during spring, and mammoni which ripen in autumn. In all three of them, the Blastophaga wasps develop synchronously with the syconia. Many of the pollen-carrying female wasps emerging from mature syconia do not land only on new caprifig syconia, but are attracted also to the numerous young female syconia borne at this time by true figs (Galil and Neeman 1977). They enter the syconia through the orifice, become trapped in them, bring about pollination, and perish. Since the wasps are unable to insert their eggs into the long-styled female flowers, the maturing ‘fruits’ of the true fig do not harbour Blastophaga larvae even after having been visited by the insects. The cultivator growing Smyrna-type figs makes use of the symbiosis between the fig and the figwasp by artificial pollination known as caprification. Twigs with mature profichi are collected from wild caprifig trees in early summer and suspended on the true fig trees in the plantation. The pollinating wasps are thus brought near to the female syconia. Caprification is an ancient procedure. It was practised in Greek and Roman times (White 1970); probably even earlier. Carbonized small seed constitute the bulk of fig tree remains retrieved from archaeological excavation. Some sites have also yielded charred dry syconia. The archaeological record of figs is sparse. It is likely to be biased since the ‘seeds’ of F. carica are small and may be overlooked.

Wild ancestry The domesticated fig tree shows a close morphological resemblance, striking similarities in climatic requirements, and tight genetic interconnections with an aggregate of wild and weedy forms that are widely distributed over the Mediterranean basin (Map 17). Botanists (Tutin 1993; Browicz 1986; Zohary 1973, p. 631) regard these spontaneous figs as the wild progenitor of the cultivated fruit trees and place them within F. carica L. Wild populations of figs (Fig. 36) grow mainly in the low altitudes of the Mediterranean maquis and garrigue formations.

They occupy rock crevices, gorges, streamsides, and similar primary habitats. They are often complemented by a wide range of feral types occupying secondary, manmade habitats such as edges of plantations, terrace walls in cultivation, ruins, collapsed cisterns, cave entrances, etc. Frequently these ‘weedy’ types seem to have been derived from seed produced by local domesticated clones that were pollinated by the adjacent wild-growing caprifigs (Zohary and Spiegel-Roy 1975; Flaishman et al. 2008) Wild females usually bear relatively small, barely edible sycons. The Mediterranean wild-feral-domesticated F. carica species-complex is closely related also to a group of non-Mediterranean, wild, deciduous Ficus taxa distributed south and east of the Mediterranean region and adapted to semi-arid and higher temperature environments (Warburg, 1905; Zohary 1973, p. 630; Browicz 1986). Taxonomically, all of them form a single natural group (series Carica in Section Eusyce) within the genus Ficus. The latter is an enormous genus comprising some 800 species distributed mostly in the tropics. Tall, large figs grow in the lower zone of the mesic, deciduous forests of the Colchic (Black Sea) district of northern Turkey and the Hyrcanic (south Caspian Sea) district of Iran and adjacent Caucasia. These forest types intergrade with the typical Mediterranean F. carica. Most authors place these mesic, wild forms within F. carica L. However, several Russian botanists (see, for example, Zhukovsky 1964), treat them as two independent species: F. colchica Grossh. and F. hyrcanica Grossh. Other members of the series Carica are warmclimate, xeric, shrubby types distributed outside the traditional area of fig cultivation: F. johannis Boiss. [syn. F. geraniifolia Miq.] in the Zagros Mountains and in southern Iran; F. virgata Roxb. in Afghanistan; F. pseudosycomorus Decne. in south Jordan, the Israeli Negev, Sinai, and Egypt; and F. palmata Forssk. in Yemen, Somalia, and Ethiopia. Some of these wild figs are interconnected by intermediate forms, and taxonomically should perhaps be considered only as eco-geographic subspecies. But none of them, with the exception of the tall, Colchic forest type, have established noteworthy contacts with the F. carica crop complex.

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Map 17 Geographical distribution of wild fig, Ficus carica. The distribution in the eastern half of the map is based mainly on data from Browicz 1986.

Several of these non-Mediterranean wild figs were genetically tested (Storey and Condit 1969) and found to have a chromosome complement identical to that of the domesticated fig (2n = 2x =26 chromosomes). All members of series Carica seem to be fully inter-fertile with the crop complex but adapted to different ecological regimes. The close affinities between the members of the Carica series are also indicated by the behaviour of the symbiotic wasp Blastophaga psenes L. At least in experimental plots, this insect moves freely between the Mediterranean and non-Mediterranean types. Due to the complex situation in fig reproductive biology, attempts to reach molecular identification and classification of fig cultivars and their wild relatives in the last decade or so were not yet fully success (see Flaishman et al. 2008; Aradhya et al. 2010).

Archaeological evidence The oldest known fig pips came from ca. 800,000 BP Mousterian Gesher Benot Ya’akov, Israel (Melamed et al. 2011). Later, charred fig pips were retrieved from numerous Early Neolithic sites in south-west Asia (Miller 1991, Table 2). The earliest Neolithic finds came from ca. 11,700–10,550 cal BP PPNA Netiv Hagdud (Kislev 1997) and ca. 11,400–11,200 Gilgal (Kislev et al. 2006a), Israel (where several

fruits were found as well). Kislev, Hartmann and Bar-Yosef (2006a), even suggest these find represent first domestication (Lev-Yadun et al. 2006, and see discussion in Kislev et al. 2006b). Additional Neolithic finds include ca. 10,500–10,200 cal BP Aceramic Neolithic Tell Aswad, Syria (van Zeist and Bakker-Heeres 1985), ca. 8,150–7,850 cal BP Nahal Zehora II, Israel (Kislev and Hartmann forthcoming) and ca. 9,300–8,500 BP PPNB ‘Ain Ghazal, Jordan (Rollefson et al. 1985). They continue to appear in this area during later Neolithic times, such as ca. 8,000–7,500 BP PPNC Atlit-Yam (Kislev et al. 2004) and the first half of the seventh millennium BP Ceramic Neolithic Jericho (Hopf 1983), Israel. In addition, fig pips were discovered in ca. 8,650–8,400 cal BP Aceramic Neolithic Knossos, Crete (Sarpaki 2009), and ceramic Neolithic Sesklo (Kroll 1981a) and Toumba Balomenou (Sarpaki 1995), Greece. Fig remains were found in some west Mediterranean early sites such as middle Neolithic Grotta dell’ Uzzo, Sicily (Costantini 1989) and Bronze Age Valeggio, north Italy (Villaret-von Rochow 1958). It is impossible to distinguish between pips of wild and cultivated figs; the excavated material can be interpreted as either. However, we are inclined to regard all these finds as representing collections from the wild. Most likely, the Carica fig was introduced into cultivation at the

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same time as its two horticultural companions: olive and grapevine. As to later sites, carbonized pips of figs were uncovered in Chalcolithic Tell Shuna North, Tell Abu Hamid, and Tuleilat Ghassul (Neef 1990), in Chalcolithic and ca. 3,500–3,300 cal BP Bronze Age Jericho (Hopf 1983), in Early Bronze Age Bab edhDhra (McCreery 1979) in the Dead Sea basin and ca. 3,450–3,350 cal BP Hala Sultan Tekke, Cyprus (Hjelmqvist 1979b). Very small pips, as well as several whole fruits, were retrieved from late Neolithic and from younger strata of fifth millennium BP Lerna Greece (Hopf 1961b). Cuneiform sources indicate that figs were grown in Mesopotamia from the second half of the fifth millennium BP on (Postgate 1987). In Egypt, the earliest archaeobotanical record of figs is from the ca. 5,450–5,650 cal. BP Maadi, and Pre-dynastic Tell elFara’in (Buto), the Nile Delta (Thanheiser 1991), and a beautiful drawing of baboons harvesting figs in twelfth dynasty Khnumhotop III’s grave (ca. 3,900 BP) in Beni Hasan (Darby et al. 1977). Contemporary indications of fig cultivation are available also from Syria (Stager 1985). They show that since the Bronze Age, figs accompanied olives and grapes as main horticultural elements of the rain-dependent agriculture in the Mediterranean basin. When the archaeobotanical data and the available information extracted from living plants are combined, one is led to the conclusion that in this fruit crop too, the earliest signs of cultivation appear in the eastern part of the Mediterranean basin and in south-west Asia. This territory harbours the closest wild relatives of the cultivated fig. In other words, the wild F. carica forms of the Levant, southern Turkey, and the Aegean belt can be regarded as the most likely ancestral wild stock from which early fig domesticates were derived. In summary, the main changes in this fruit crop under domestication were the shift to vegetative propagation of female clones as follows: (i) the increase of the size, amount of flesh, and sugar content of the syconia; (ii) the introduction of artificial pollination and caprification; and (iii) the effective selection for parthenocarpy where pollination is not necessary for fruit set. Such developments were already part of fig horticulture in classical times.

Sycamore fig: Ficus sycomorus The cultivation of the sycamore fig, Ficus sycomorus L. (Moraceae), has been almost exclusively an Egyptian speciality. Compared with the common fig, F. carica, it is a much taller and larger tree, but it produces smaller, inferior syconia. Since early dynastic times, ca. 6,000 years BP, the sycamore fig was (and still is) a commonly used fruit crop and a valued timber source in the lower Nile Valley (Täckholm 1976). Outside Egypt, the sycamore fig was less esteemed, though it was introduced into the warmer parts of Israel and to a lesser extent to several other locations on the shores of the Mediterranean Sea (e.g. Lebanon, Cyprus, Tunisia). As with other members of the genus Ficus, the reproduction of the wild trees is from seed, and seed setting depends on pollination by a specific symbiotic wasp, Ceratosolen arabicus Mayr. In contrast, cultivation depends on clonal propagation (rooting of twigs), and fruit production no longer depends on the natural pollinator. Some clones are parthenocarpic, (set fruit without fertilization). In others, fruit maturation is artificially induced by gashing the surface of the young syconia. This is an old tradition in Egypt and in Israel (Galil et al. 1976). Wild F. sycomorus is widely distributed in eastern Africa from Sudan to South Africa, with an extension to Yemen. The tall, big trees, growing near streams and in beds of ephemeral watercourses, constitute a conspicuous component of savannah landscapes. Throughout this area, F. sycomorus reproduces sexually. As previously mentioned, in this fig a specific symbiotic Ceratosolen wasp brings about pollination. As far as we know (Galil et al. 1976), spontaneously reproducing sycamore figs, as well as their pollinating wasp, do not occur today in Egypt, but they still thrive in nearby Sudan.

Archaeological evidence Remains of F. sycomorus start to appear in Egypt in pre-dynastic times. The earliest records come from ca. 6,000–5,500 BP Amratian sites, and from ca. 4,600–4,400 BP El Omari (Wetterstrom 1993, 1998). Shortly after, sycamore fig remains appear in quantity in Egyptian excavations from the start of the

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fifth millennium BP onward (Galil et al. 1976; Germer 1985; Murray 2000b). The fruit and the wood, and sometimes even the twigs, are richly represented in the tombs of the Early, Middle, and Late Kingdoms. In some cases, the parched sycons bear characteristic gashing marks indicating that this art was practised in Egypt already in ancient times. There is no doubt that Egypt was the principal area of sycamore fig development. In spite of the absence of spontaneous trees and their pollinating wasp in Egypt at present, it seems that F. sycomorus was brought into domestication in Egypt (Galil et al. 1976).

Date palm: Phoenix dactylifera The date palm, Phoenix dactylifera L. (Palmae), ranks among of the first fruit trees that were brought into domestication in the Old World. It was already part of south-west Asian food production in the Chalcolithic period (Zohary and Spiegel-Roy 1975). Compared with the three other classic fruit trees (olive, grapevine, fig), the date palm requires a much warmer and dry-summer climate. High temperatures, rainless summers, and very low humidity are particularly important for fruit setting and fruit ripening. Date horticulture is therefore centred in the deserts south of the Mediterranean Sea and in the southern fringe of south-west Asia (from south Iran to the Atlantic coast of north-west Africa). This is an almost rainless belt, in which date horticulture depends on mild winters, hot and dry summers, and a steady water supply, either by the presence of a high level of ground water or by artificial irrigation. Date palms can withstand considerable salinity and they grow well even when watered by brackish water. Arab folklore has cleverly summed up the ecology of the date palm: ‘Its feet are in the water and its head is in the fire of the sky’. Date palms are very productive and the annual fruit yield may be as high as 100–200 kg per tree. All over these warm Arabian and Saharan deserts, the sugarrich fruits (some 60–70% sugar content) serve as an important staple food for local people. Other parts of the palm tree are also intensely used. In addition, the worldwide date-palm production has grown from 1.75 million metric tons in 1962 to 7.2 million

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metric tons in 2009 (http://faostat.fao.org). At present, the largest producer of dates is Egypt (Chao and Krueger 2007). The trunks are split longitudinally into halves or quarters, and serve as beams for building, the leaves are usedwidely for roofing, matting, and basketry, and the fibres of the bark are used to prepare ropes. Date palms start to bear fruits four to five years after planting, and reach full fruit production at eight to ten years. For details on date palm cultivation and on its reproductive biology, consult Nixon (1951), Dowson (1982), Reuveni (1986), Chao and Krueger (2007). Phoenix dactylifera is a diploid (2n = 2x = 36 chromosomes) dioecious palm characterized by its tall trunk (up to 25 m), its large, leathery, feather-shaped leaves, and its ability (when young) to produce basal suckers. The vegetative propagation of clones, practised by the farmer, depends on these suckers. Wild populations, as well as seedlings derived from domesticated clones, consist of an equal proportion of female and male individuals. This is brought about by a single gene, or block of genes, determining sexuality. The male morph is heterozygous and the female homozygous recessive. Date palms are wind-pollinated. In domesticated palm-groves, artificial pollination is applied by the farmer, and the fraction of male individuals required for effective pollination is reduced to one male per twentyfive to fifty female trees. This tradition is very old (Schiel 1913; Pruessner 1920). Apparently it was practiced in Mesopotamia already in Hammurabi’s time (1,792–1,750 BC).

Wild ancestry The wild stock from which the domesticated date palm could have been derived has been satisfactorily identified (Zohary and Spiegel-Roy 1975). The crop is closely related to a variable aggregate of wild and feral palms distributed over the hot and dry southern parts of south-west Asia as well as the north-eastern Saharan and North Arabian deserts (Map 18). These spontaneous dates show close morphological similarities and parallel climatic requirements with the domesticated clones. In addition, they are fully inter-fertile with the cultivars and are interconnected with them by occasional hybridization. Botanists place these wild-growing date palm

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populations within the species complex of P. dactylifera L. The wild-growing forms produce basal suckers just as the domesticated varieties, but differ from the domesticated clones by their smaller fruits. These wild dates contain relatively little pulp and are frequently unpalatable (or even indigestible). Thus, in the date palm, domestication has led to the increase in fruit size and pulp quality. Wild and domesticated date palms differ in their mode of reproduction. Sexual reproduction is the rule in wild stands, while domestication has brought a shift to vegetative propagation—to the ‘fixation’ of desired highly heterozygous female clones. P. dactylifera was well suited for this shift since it produces easily transferable suckers. Spontaneously growing dates occur in almost the entire area of date cultivation (Map 18) and frequently represent secondary escapees. However, in some areas in south-west Asia and very probably also in the north-eastern Sahara, Arabia, and Baluchistan, dates are genuinely wild and occupy primary niches. Prominent among these locations are lowland Khuzistan—the southern base of the

Zagros Range facing the Persian Gulf—and the southern part of the Dead Sea basin (Zohary and Spiegel-Roy 1975). In these exceptionally hot, dry territories, wild-type dactylifera palms, with their characteristic small and mostly inedible fruits, thrive in gorges, wet rocky escarpments, seepage areas in wadi beds, and brackish springs, where they constitute a conspicuous element in the vegetation. In P. dactylifera we are therefore faced with a variable complex of wild forms, segregating escapees, and domesticated clones, which are all genetically interconnected by occasional hybridization. It is often difficult to decide whether non-domesticated material is genuinely wild, or whether it represents weedy forms or secondary seedlings derived from domesticated clones. However, it is impossible to delimit the pre-agriculture distribution of the wild date. There can be little doubt that the wild dactylifera forms are indigenous to the hot and dry parts of south and west Asia. Date palms thrived in these territories long before the initiation of agriculture. This is attested by the finds of charred pieces of P. dactylifera wood in 23,000 BP retrieved from Ohalo

0 200 400 miles 0 200 400 km Map 18 Geographical distribution of wild and feral forms of date palm, Phoenix dactylifera (based on Fischer 1881). In south Arabia, Sudan, Eritrea, Somalia, and Senegal they sometimes cross with the native P. reclinata; and in the Indus basin with P. sylvestris.

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Fig. 37 Carbonized stones of date palm, Phoenix dactylifera, Chalcolithic Tuleilat Ghassul, Jordan (Zohary and Spiegel-Roy 1975).

II (Liphschitz and Nadel 1997), and the presence of numerous date-pollen grains in Mousterian and Baradostian strata in Shanidar Cave in the Zagros foothills (Solecki 1975). Phoenix dactylifera is the main south-west Asian wild representative of its genus that comprises some thirteen species distributed over Africa and south Asia (Jones 1995; Barrow 1998). The only other wild date that occurs in the east Mediterranean basin is P. theophrastii Greuter, a narrow endemic date palm confined to Crete and several locations in coastal south-west Turkey (Barrow 1998). Several species of Phoenix have been tested cytogenetically. All show a constant chromosome complement (2n = 2x = 36 chromosomes) and are fully inter-fertile with one another. As in the case of Vitis, species in Phoenix are reproductively isolated from one another not by inter-sterility but by geographical and ecological barriers. Two additional wild Phoenix species grow on the fringe of traditional date cultivation in the Old World and, in these areas, they may have enriched the gene pool of the domesti-

cated fruit trees through spontaneous hybridization. In south Arabia, and in Africa south of the Sahara, date cultivation comes in contact with the bushy P. reclinata Jack., whereas in the Indus Valley contact is achieved with the tall, non-suckering, more tropical P. sylvestris Roxb. In both territories, hybridization between the cultivated date and the native wild species has been detected (Munier 1973). Recently, Rhouma et al. (2008) employed random amplified microsatellite polymorphism markers (RAMPOs) among thirty date-palm cultivars and ten male trees to assess the genetic diversity and phylogenic relationships in date palms. Their results support the Mesopotamian origin of the date-palm domestication.

Archaeological evidence Archaeological sites with date palm remains are not abundant so far, but this may reflect the fact that most excavations in south-west Asia and the

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Mediterranean basin were conducted in areas too cold for date-palm cultivation. The few stones of date palm reported from Egypt, Israel, Iran, and Pakistan, dated eighth and seventh millennia BP, probably represent material collected from the wild. The earliest remains of what seem to be domesticated dates have been found by Seton Lloyd in the ca 6,000 BP Ubaidian horizon at Eridu, Lower Mesopotamia (Gillett 1981). R.J. Braidwood (pers. comm.) remarked that ‘buckets of date stones’ were uncovered in this site. In Jordan, charred kernels (Fig. 37) were uncovered in the ca. 6,800–5,800 cal BP classic Chalcolithic Tuleilat Ghassul in the Jordan valley (Zohary and Spiegel-Roy 1975). Date kernels were also retrieved from ca. 5.500–5,200 BP Chalcolithic of the ‘Cave of the Treasure’ in Nahal Mishmar, Israel (Bar-Adon 1980), and a single date kernel is available from Early Bronze Age Jericho (Hopf 1983). Date kernels were also found among offerings deposited in a tomb in the ‘Royal Cemetery’ at Ur, lower Mesopotamia and dated to the late fifth millennium BP (Ellison et al. 1978). Fruits and kernels from numerous archaeological sites from the early sixth millennium BP across eastern Arabia indicate its important role ever since the Early Bronze Age (Cleuziou and Costantini 1982; Nesbitt 1993; Méry and Tengberg 2009; Beech 2001, 2003). Indeed, the occurrence of these finds near current and past habitats of wild date, emphasize the need to establish their domestication status, beyond the sheer quantity of kernels in some of the finds. From the Bronze Age on, date cultivation seems well established in the warmer sections of south-west Asia. Dates are frequently mentioned in Sumerian and Akkadian cuneiform sources from the second half of the fifth millennium BP onwards (Postgate 1987; Nesbitt 1993). They are depicted also on seal engravings. The written documentation shows that by the Late Uruk period (ca. 5,400–5,100 BP) the cities of Sumer possessed flourishing date plantations. Remains of dates appear rarely in Pre-dynastic Egypt, such as Hierakonpolis and El Omari, and prior to the Middle Kingdom much of the date remains consist of leaves, fibre, and wood, rather than fruits (Murray 2000b). Täckholm (1976) and Murray (2000b) stresses that most finds are from the New Kingdom and post-Pharaonic periods. Täckholm (1976) noted that in the New Kingdom

remains of date fruits occur ‘in almost every second tomb; in all forms, all sizes, and all colours’. Since the second half of the fifth millennium BP, datepalm cultivation seems to have been practised also in the dry and warm parts of Baluchistan and in adjacent parts of the Indian subcontinent (Costantini and Costantini-Biasini 1985). In classical times, dates appear to be an important food element in lower Mesopotamia, Oman, the lower Jordan Valley, the Nile Valley, and the desert oases of North Africa, Arabia, and Baluchistan. The available archaeological data, as well as the information on the living date palm, seem to focus on south-west Asia as the initial place of P. dactylifera domestication. This area which furnished the earliest indications on date palm cultivation also harbours wild forms from which the first cultivated clones could have been obtained. The date palm was probably brought into cultivation somewhere in the lower Mesopotamian basin, or in some oases in the southern fringe of the Fertile Crescent.

Pomegranate: Punica granatum The pomegranate, Punica granatum L. (Punicaceae), is an appreciated minor crop in traditional Mediterranean horticulture. It is a deciduous bush or small tree with conspicuous red flowers and large (6–12 cm across) fruits characterized by a leathery rind, persistent crown-like calyx, and numerous seed covered with a juicy flesh which can be eaten fresh, or whose juice can be extracted, and also be fermented into wine. The wild ancestor of the domesticated pomegranate has been satisfactorily identified (Zohary and Spiegel-Roy 1975). Wild forms of P. granatum grow in masses in the south Caspian belt, in northeastern Turkey, and in Albania and Montenegro. Domestication brought about an increase in fruit size and a shift from sexual reproduction to clonal propagation. Desired genotypes are easily maintained by rooting of winter-dormant suckers. Cultivars set fruit by self-pollination.

Archaeological evidence The pomegranate seems to belong to the early ensemble of domesticated fruit trees in the Old

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World. In addition to supplying fresh fruit, they were also considered to be a symbol of fertility. The earliest carbonized pips and fragments of pomegranate rind have been obtained from Early Bronze Age Jericho (Hopf 1983), Arad (Hopf 1978c), and in the burned Palace’s kitchen in ca. 4,900 cal. BP Tell es-Sa’idiyeh (ancient Zaretan), Jordan (Cartwright 2003, and pers. comm.). They also appear in Late Bronze Age ca. 3,450–3,350 cal BP Hala Sultan Tekke, Cyprus (Hjelmqvist 1979a), and Tiryns, Greece (Kroll 1982). An interesting find is the numerous seeds and fruit fragments in a large jar, and hundreds of seeds across the site, from the late twentyfourth century BP Late Bronze Age Uluburun shipwreck (Haldane 1993). As Ward (2003) indicates, this is an indication of the pomegranate’s status as a luxury item in Bronze Age eastern Mediterranean culture. Pomegranates are recorded in several ancient Mesopotamian cuneiform sources—from the second half of the fifth millennium BP on (Postgate 1987). In fact, pomegranate is depicted in artefacts and paintings before its appearance as plant remains, first occurring in Mesopotamia (see details and discussion in Ward 2003). In Egypt, Schweinfurth (1891) identified pomegranates among the vegetable remains of the twelfth dynasty (ca. 1990–1800 BC) Drah Abu el Naga, Egypt. A large, dry pomegranate was found in the tomb of Djehuty, the butler of Queen Hatshepsut, dating from about 1470 BC (Hepper 1990). Pomegranates do not occur in a wild state in south Levant. Therefore, the occurrence of Punica remains in Early Bronze Age in Israel and Jordan suggest well established cultivation.

Apple: Malus domestica The domestic European apple, Malus domestica Borkh. (= M. pumila Mill.) of the rose family (Rosaceae), is the most important (63 million tons for 2008) fruit crop of the temperate, cooler parts of the Old World (Way et al. 1991; Watkins 1995; http://faostat.fao.org). Since Greek and Roman times, apples have been extensively grown in the temperate parts of Europe, south-west Asia, and probably also in the Caucasus and in central Asia. They thrive in regions where winters are sufficiently cold to provide this fruit crop with a chilling phase

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necessary for breaking bud-dormancy. The fruit is a pome, a ‘false fruit’, the greater part of which is formed not by the ovary but by the receptacle of the flower. In the core of the pomes of most cultivars there are five leathery chambers or loculi, each normally containing two seeds. Apples under domestication are very variable. Some 2000 distinct cultivars have already been described. They vary in fruit size (3–12 cm in diameter), colour (red, yellow, green), shape, and in the texture and taste of the pulp (sweet to acid). The majority of present-day cultivars belong to the group of dessert apples that bear relatively sweet fruits that are consumed fresh, or are used to prepare compotes or pies. Cooking apple varieties produce harder and often larger pomes. Other cultivars are cider apples, selected for production of vintage quality cider. Less fancy types of this alcoholic beverage can be prepared by fermenting ordinary dessert apples or even wild ‘crab apples’. Finally, some apples are grown for their ornamental features, while other ones serve as rootstocks on which superior scions are grafted.

Wild ancestry The apple, Malus Mill., contains some fifteen indigenous, ‘primary’ wild species. Most of them are self-incompatible, contain 2n = 2x = 34 chromosomes, can be crossed with one another, and their interspecific hybrids are fertile or at least partially fertile (Korban 1986; Janick et al. 2003; Richards et al. 2009). However, some cultivars and wild forms are triploid (2n = 3x = 51 chromosomes) and a few are tetraploid (2n = 4x = 68 chromosomes). The domesticated apple varieties (usually referred to as M. domestica Borkh. or as M. pumila Mill.) are closely related to a variable group of wild ‘crab apples’ (series Pumilae in section Malus of the genus Malus (Rehder 1940; Way et al. 1991). It is a variable domestic crop species complex that contains also a large number of ‘secondary’ inter-specific hybrids, introgressive hybridization products, and feral populations. These wild apples (Plate 14) are widely distributed over the temperate forest zone of Europe, south-west and central Asia, and in Siberia. Closest

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to the crop (and freely crossing with it) are the European, Caucasian, and Siberian wild crab apple forms, which are placed by some apple taxonomists in M. sylvestris (L.) Miller. In other taxonomic treatments, the western populations of these crab apples are kept as M. sylvestris and M. orientalis, while Siberian forms are very variable and regarded as a third subspecies, M. sieversii (Ledeb.) M. Roem. (Janick et al. 2003). Taxonomically, Malus is relatively a large genus, containing several sections, subsections and series. The crop (together with its close wild relatives) is placed in section Malus subsection Malus series Pumilae, of the genus Malus (Rehder 1940; Way et al. 1991, p. 30; Watkins 1995; Janick et al. 2003). This is the largest section in this genus. Its members are widely distributed over the temperate and cool forest areas of Europe and Asia from the Atlantic coast of Europe to China, and expand to North America. As stressed by Korban (1986), by Way et al. (1991) and by Watkins (1995) most of the wild species, as well as the domestica cultivars, cross readily with one another. Their hybrids and ‘hybrid swarms’ are fertile or at least partly fertile. This is particularly the case with the domesticated apple varieties. If they are introduced and planted close to their wild relatives, they readily cross with them. Before apple domestication, wild apple populations were apparently isolated from one another mainly by geographic and ecological reproductive barriers. However, this type of isolation broke down when apple-culture moved in and tied these variable taxa together. Wild-growing crab apples reproduce from seed, and are self-incompatible. They are characterized by small fruits (1.5–3.0 cm in diameter), and are very variable in their shape, colour, and taste. The majority of plant taxonomists rank all these forms as eco-subspecific units of the European wild ‘crab apple’ M. sylvestris (L.) Mill. (Map 19). However, because they show a marked differentiation from eco-geographical races, some taxonomists regard them as independent, vicarious species. Forms growing in Europe are usually referred to as M. sylvestris (L.) Mill. subsp. sylvestris. Forms growing in Anatolia, the Caucasus and Transcaucasus, and northern Iran are regarded (Browicz 1972) as a distinct regional subspecies, M. sylvestris subsp. orien-

talis (Uglitzkich) Browicz. Further east M. sylvestris is replaced in central Asia and Siberia, by a group of central Asiatic wild crab apples that are either grouped in M. sieversii (Ledeb.) M. Roem. or split even further by Russian botanists who recognize several additional local species: M. kirghizorum A. & Fed. and M. turkmenorum Juz. & M. Pop. The eastern Siberian and north Chinese forms are placed in M. prunifolia (Willd.) Borkh. Variation in apples is further complicated by the fact that in numerous places in Europe, south-west Asia, the Caucasus, and in central Asia, apple cultivation has been practised in areas also supporting wild apple populations. Furthermore, in some regions—for example, in Anatolia and the Balkans—wild apples were commonly used, as in situ stock material for grafting. Under such conditions, spontaneous hybridization between tame and wild apples occurs quite frequently, resulting in the formation of swarms of secondary hybrid derivatives. Recently, crossing of the European apples has further complicated variation patterns in apples with numerous wild and cultivated forms introduced from other geographical origins (particularly from central and Eastern Asia). This development has already produced a whole array of new hybrid combinations. Based on morphological traits and several genetic tools of species relationships, most researchers (e.g. Wagner and Weeden 2000; Forte et al. 2002; Harris and Juniper. 2002; Richards et al. 2009), support the central Asian apple M. sieversii as the ancestor of domesticated apple. These studies point to the wild M. sieversii forests of Tian Shan Mountains of central Asia as probable center of origin. In particular, the recent full genome sequencing of Velasco et al. (2010) validates M. sieversii as the progenitor of the cultivated apple. It also rejected the view of Coart et al. (2006) who concluded that the European crab apple M. sylvestris is the ancestor of domesticated apples. Taxonomically, if M. sieversii is the wild progenitor of the domesticated crop, domesticated apple should be regarded as its subspecies.

Archaeological evidence Apples no doubt were collected from the wild long before their domestication and before the

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0 100 200 miles 0 200 400 km

Map 19 Geographical distribution of the European crab apple, Malus sylvestris (including subsp. orientalis) (based mainly on Browicz 1972, and pers. comm.). Other wild apple forms, closely related to the domesticated apple, are common in central and east Asia (Forsline et al. 2003).

establishment of vegetatively propagated clones. Numerous carbonized remains of small apples (2–27 mm in diameter), often cut into halves for parching, were uncovered in Neolithic and Bronze Age Switzerland (Schweingruber 1979). These remains correspond morphologically very well to the wild crab apples that grow today it that area. Similar remains of wild, small apples were retrieved in numerous Neolithic and Bronze Age sites in Europe; for example in former Yugoslavia, Hungary, Poland, Czechoslovakia, Austria, Germany, and Denmark (for an enumeration of finds, see Hopf 1973b; SchultzeMotel 1968–1994; Kroll 1995–2000, as well as the online databases http://www.cuminum.de/archaeobotany/database/; http://www.archaeobotany.de/ database.html). Small apples, cut transversely into halves, threaded and dried on strings, were found among offerings deposited in a tomb in the ‘Royal Cemetry’ at Ur, lower Mesopotamia, and dated to the Sargonid period, late fifth millennium BP (Ellison et al. 1978).

This discovery confirmed descriptions of strings of apples in cuneiform texts from the same time. Since apples do not grow wild in warm and dry Mesopotamia, such small apples could represent a long-distance import of wild apples or early attempts of apple cultivation. Because Malus very rarely lends itself to simple vegetative propagation, apple cultivation in these early times would have been necessarily based on seed planting; in other words, on handling of variable segregating individuals resembling ‘crab apples’. This is indeed what the small, stringed apples look like. Yet even such low-grade apples seem to have been appreciated in ancient Babylonia, and were used for offering. A similar early indication of apple cultivation comes from ca. tenth century BC Iron Age Kadesh Barne’a on the border between the Negev and Sinai. Here several dozens of well-preserved charred fruits have been discovered (Rudolph Cohen, pers. comm.). Wild apples do not occur in this area and their southernmost limit today is Turkey. These

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remains, therefore, strongly suggest that apples were cultivated in the oasis of Kadesh Barne’a. Whatever the nature of the Ur and Kadesh Barne’a finds, one fact is clear: the apple did not evolve into a major fruit crop in Bronze Age times. While the cultivation of the olive, the grapevine, and the date palm accelerated dramatically in the fifth and fourth millennia BP, there are no signs of a similar development in the apple. This tree seems to have obtained its significant role in the Old World horticulture only in classical times. This development is apparently linked to the introduction of grafting. The Greeks were already familiar with the art of grafting (White 1970, p. 248) and Theophrastus writes about the maintenance of apple clones by this sophisticated method of vegetative propagation. In conclusion, we still know very little about the time and place of apple domestication except that in classical times apples were already extensively grown in the Old World. Wild apples are widely distributed over Europe and west Asia, where they have diverged into numerous types and closely related species that still retain full inter-fertility. Apple cultivation is superimposed on this wild background. It is therefore futile to try to delimit the area of initial domestication on the basis of the evidence available from the living plants. Most (although not all) genetic evidence indicate that the apple was domesticated in the Tian Shan Mountains of central Asia, and reached Europe with remarkably little later hybridization. Even so, exceptional Malus individuals may have been picked up not once and in a single place, but many times and in several areas. Furthermore, many cultivars are hybridization products combining genes from several distinct geographical sources. Nevertheless, most of the traditional varieties in Europe and south-west Asia have their closest morphological affinities with the wild M. sylvestris and this undoubtedly is the principal, but not the only, wild source from which the old cultigens in these regions have evolved. The archaeological finds make it clear that apples have been extensively collected from the wild during Neolithic and Bronze Age times. In the Bronze Age and the Iron Age they may have been occasionally planted from seed (as a luxury fruit). However, the first signs of substantial apple cultivation appear

much later than those of olive, date-palm, fig, or grapevine horticulture. The introduction of grafting is apparently a key element in this development and actually defines domestication. Because apple cultivation depends almost entirely on this method of vegetative propagation, it could only have been practised after this technology became known in south-west Asia and Europe. Where and when grafting was invented is not clear, although its initiation was probably not in the area of Mediterranean horticulture (see p. 115).

Pear: Pyrus communis The domesticated pear of Europe and west Asia Pyrus communis L. [= P. domestica Med.] (Rosaceae), is second to the apple in its contribution to fruit production in the cool and temperate parts of Old World agriculture (Watkins 1986). Frequently, pears are close companions of apples, and both fruit crops thrive in similar environments. Pears, too, require sufficient winter chilling to ensure normal flowering and fruit setting. As in apples, the fruits are pomes in which the larger part is formed by the receptacle of the flower and only the inner ‘core’ containing the seed, develops from the ovary. Usually the pomes have the distinctive pyriform shape of Pyrus and their pulp contains stone cells that give pears their characteristic gritty texture. Domesticated European pears are extremely variable. About one thousand distinct cultivars are known by their varietal names. Like most other fruit trees of the rose family, pears are self-incompatible (Zielinski et al. 1965). Wild pears are diploid (2n = 2x = 34 chromosomes), and the majority of domestic clones are also diploid. However, some of the latter are triploid (2n = 3x = 51 chromosomes) or tetraploid (2n = 4x = 68 chromosomes). Pears segregate widely when grown from seed, and most of the progeny is worthless. Like apples, the maintenance of desired clones depends almost entirely on grafting.

Wild ancestry The genus Pyrus L. comprises some thirty wild species distributed over Europe and Asia from the Atlantic shore of west Europe to China (Rehder

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0 100 200 miles 0 200 400 km

Map 20 Geographical distribution of wild forms of the European pear, Pyrus communis (based on Browicz 1992; additional information kindly provided by this author). A few other wild pear forms, closely related to the domesticated pear, are growing throughout Asia (Dzhangaliev et al. 2003).

1940; Browicz 1993). Different wild species of pears were domesticated in Europe, and independently in east Asia. The domesticated pear of Europe (P. communis) is closely related to a variable group of wild pears distributed over the temperate parts of Europe, northern Turkey and the Caucasus (Map 20). The wild populations that thrive in temperate Europe are frequently called P. pyraster Burgstd. Those growing in the eastern parts of Asia are referred to as P. caucasica Fed. Because of their tight morphological and genetic affinities and full inter-fertility with the crop, these wild pears are now considered as the two eco-geographic wild subspecies of the crop’s complex from which the European domestic pear P. communis could have been derived. Most of these wild pears are rather spinescent and bear very gritty small fruits (1.5– 3.0 cm in diameter). In summary, all wild populations are diploid (2n = 2x = 34 chromosomes) self-incompatible trees, and all are inter-fertile with the European domesticated crop.

Spontaneous crosses between these wild pears and domesticated clones are quite frequent. In numerous localities in Europe and western Asia, pyraster or caucasica wild forms and communis cultivars are interconnected by feral forms and hybridization derivatives that thrive best at edges of cultivation and in clearings of forests adjacent to cultivation. Armenia and the Caucasus are extremely rich in wild pear forms. Because of their tight morphological and genetic affinities with the crop, Browicz (1992, 1993) in his taxonomic treatment of Pyrus, placed pyraster and caucasica pears as wild subspecies, in P. communis. Several other wild pears that grow in the general area of traditional pear cultivation are inter-fertile with the domesticated crop. Some or even all of these species have enriched the genetic variation of domesticated pears through hybridization and introgression (Rubtzov 1944; Watkins 1986). Prominent among these wild pears are P. spinosa

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Forssk. [= P. amygdaliformis Vill.], native to west Turkey, the Aegean basin, and the south Balkans; P. elaeagrifolia Pallas distributed over Turkey and east Bulgaria; P. salicifolia Pallas in Caucasia and adjacent areas in Turkey; the more arid P. syriaca Boiss. of the Fertile Crescent (Plates 15 and 16); and its closely related P. korshinskyi Litw. in central Asia (for details on the distribution of these species see Browicz 1982, pp. 49–52 and maps 79–83.) In earlier times, wild pears were frequently grafted in situ. This practice is still common in Anatolia, where farmers frequently spare individuals of wild P. communis, P. syriaca, P. spinosa, and P. elaeagrifolia at the edges of cultivation and inside grain crop fields. These trees are grafted with domesticated clones. Often only one or a few main branches bear scions; the others are left as they are. In such situations, cross-pollination between the two types is almost inevitable. Finally, it should be pointed out that several wild pears native to northern China and eastern Siberia were apparently brought into domestication independently and gave rise to the Far East pear cultivars (Watkins, 1986). These include: (i) the Chinese sand pear, P. pyrifolia (Burm.) Nakai, (ii) the Ussurian pear, P. ussuriensis Maxim, and (iii) the Chinese white pear P. bretschneideri Rehd. Crosses between P. communis and these three east Asian pears resulted in the selection of new commercial pear varieties and have caused further extensive genetic fusion in the genus Pyrus.

Archaeological evidence Pears were collected from the wild long before their introduction into cultivation. Charred remains of small fruits, sometimes halved, and probably dried, were found in several Neolithic and Bronze Age sites in Europe, e.g. north Italy, Switzerland, Yugoslavia, and Germany (for enumeration of sites, see Hopf 1978a; Schultze-Motel 1968–1994; Kroll 1995–2000; as well as the online databases http:// www.cuminum.de/archaeobotany/database/ ; http://www.archaeobotany.de/database.html) . Similar finds are available from late Neolithic Dimini and Bronze Age Kastanas, Greece (Kroll 1983), and from some Tripolye culture settlements in Moldavia and the Ukraine (Januševič 1978).

The archaeological evidence until now does not provide definite clues about the beginning of pear domestication. Reliable information on pear cultivation first appears in the works of the Greek and the Roman writers (Hedrick et al. 1921; White 1970). Fruit growers learned by experience that a grafted branch gave fruit true to type, whereas seedlings tended to revert to the wild state or to produce fruit of poor quality. Theophrastus (ca. 371–287 BC) describes three cultivated varieties in Greece and mentions that they were propagated by grafting since propagation by seeds resulted in inferior progeny. Cato (235–150 BC) described methods for pear cultivation and reported on six pear cultivars grown in his time. Somewhat later, Pliny (23–79 AD) recognized thirty-five cultivars. The range of fruit characters described by these authors covers many types of today’s pears. In conclusion, the patterns of variation in Old World pears closely resemble those of apples. Wild pears are widely distributed over Europe and western Asia and have diverged into numerous local ecogeographical races and species, though they retain their inter-fertility. Pear cultivation is superimposed on wild populations and hybridization between wild and tame pears has repeatedly produced secondary hybrid derivatives that tend to colonize disturbed grounds. Faced with a crop complex of such a vast distribution and extensive genetic fusion, one can scarcely determine an area of origin of pear cultivation on the basis of the living plants. Various regions in west Asia and in Europe, within the distribution range of pyraster and caucasica pears, are equal possibilities.

Plum: Prunus domestica Plums rank third to apples and pears in terms of their role in fruit production in the cooler and temperate parts of the world. They are eaten fresh, cooked, or dried as prunes. The principal cultivated plums of Europe and south-west Asia are hexaploid (2n = 6x = 48 chromosomes) Prunus domestica L. (Watkins 1986; Ramming and Coicu 1995; KörberGrohne 1996). This is a variable fruit crop in the Rosaceae, comprising two fully inter-fertile groups of cultivars: (i) the relatively small fruited damsons,

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bullaces, and greengages, P. domestica L. subsp. insititia (L.) C. K. Schneider [= P. insititia L.]; and (ii) the common European plums, P. domestica L. subsp. domestica C. K. Schneider [= P. domestica L. sensu stricto], with relatively large fruits. Also grown in this part of the world is the closely related cherry plum or myroblan, P. cerasifera Ehrh. [= P. divaricata Ledeb.], with fruits resembling those of the damsons. P. cerasifera is mainly diploid (2n = 2x = 16 chromosomes), but contains also tetraploid (2n = 4x = 32 chromosomes) and hexaploid (2n = 6x = 48 chromosomes) forms. Over most of Europe, the cherry plum is used mainly as an ornamental, or as root-stock for grafting, and cultivars planted for their fruits are rather rare (Stika and Frank 1988). However, in the Caucasus, fruit bearing cherry plums (Alyčia cultivars) constitute an important fruit crop (Hanelt 1997). In central Europe, the tetraploid (2n = 4x = 32 chromosomes) sloe, P. spinosa L., have been taken into cultivation locally. Clones of this spiny plum, with relatively larger and less astringent fruits, have been selected. They still survive (frequently in hedges) particularly in the Czech Republic, Slovakia, Austria and Armenia (Hanelt 1997; Gabrielian and Zohary 2004). Other plums, belonging to the diploid (2n = 2x = 16 chromosomes) P. salicina Lindl. have been taken into cultivation in east Asia and are known as Japanese plums. Until about 130 years ago, the European plums and the Japanese plums were separate crops, but in recent years, P. domestica cultivars constitute an important fruit crop (Hanelt 1997). In central Europe, tetraploid cultivars were crossed with the Japanese plum. Some of the modern plum cultivars are derivatives of these crosses. Plum cultivation depends almost entirely on grafting. Traditional methods of vegetative propagation (rooting of shoots, etc.) do not succeed in this fruit tree, except in the case of some suckering insititia forms. Reproduction from seed is also impractical since cultivars are highly heterozygous and their progeny segregate widely. Most of the seedlings are economically worthless. The majority of hexaploid P. domestica clones are self-compatible or at least partly self-compatible. In contrast the diploid plums are self-incompatible.

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Wild ancestry The plums comprise a distinct taxonomic group (of about two species), distributed over the temperate parts of Eurasia and north America. Some botanists follow the traditional classification and regard the plums as an independent genus (Prunus L. sensu stricto). Others, taking into consideration the fact that plums can be crossed to several other stonefruit trees, regard them only as a subgenus within an extended Prunus L. (which also includes the cherries, almonds, peaches, apricots, laurel cherries, etc.). The closest wild relatives of the hexaploid common plum are the spontaneous, somewhat spinescent, insititia-like populations of P. domestica. As argued by Werneck and Bertsch (1959), these populations suggest that P. domestica could have preceded agriculture and should be regarded as an indigenous element in middle Europe. They might be native also in the Balkans and Turkey. They bear small (2–3 cm) subglobose fruits, which are quite common in many parts of temperate Europe and Turkey. They thrive in woods, cleared hillsides, edges of cultivation, and hedges. European botanists (see, for example, Webb 1968) regard them as feral or naturalized elements. However, pre-Neolithic remains of carbonized plum stones discovered in the Upper Rhine and the Danube regions closely resemble the stones of present day. In addition, the hexaploid European plums (P. domestica subsp. domestica) and the hexaploid damsons, bullaces, and greengages (P. domestica subsp. insititia) are closely related to a variable aggregate of wild and domesticated plums grouped in P. cerasifera Ehrh. [= P. divaricata Ledeb.]. The wild forms of this aggregate are distributed over the Balkans, Turkey, the Levant, Caucasia, the south Caspian coast, and central Asia (Browicz, 1996, Map 21). They are self-incompatible, reproduce from seed and bear roundish, small (ca. 2 cm in diameter) green, purple, or dark violet fruits, which taste very much like the domesticated P. domestica plums. P. cerasifera [= P. divaricata] wild forms fall into several eco-geographical subspecies (Browicz 1996): (i) subsp. cerasifera, with glabrous leaves and twigs, grows in the more temperate northern territories; (ii)

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subsp. caspica (Kovalev & V. Ekim) Luneva [= P. caspica Kovalev & V. Ekim], with pubescent petioles, pedicels and leaves with velutinous lower surface, occur in the Caspian coast of Caucasia and adjacent Iran; (iii) subsp. ursina (Kotschy) Browicz [= P. ursina Kotschy], with short pedicels and more or less pubescent shoots and leaves, is native to the Levant and south-east Turkey; and (iv) the smaller subsp. sogdiana (Vassilez.) Cinovskis [= P. sogdiana Vassilez.] native to central Asia (Tadzhikistan, Uzbekistan, and southern Kirgizia). Chromosomally, wild P. cerasifera is not yet adequately studied. Yet, it is clear that it comprises diploid (2n = 2x = 16 chromosomes) forms (apparently the common chromosome level) as well as tetraploid (2n = 4x = 32 chromosomes) and hexaploid (2n = 6x = 48 chromosomes) chromosome races (Beridze and Kvatchadze 1981; Watkins 1986) which are morphologically very similar to their diploid relatives. Closely related to P. domestica (and to P. cerasifera) are two additional wild plums: (i) P. cocomilia Ten., a diploid (2n = 2x = 16 chromosomes) small tree, native to the southern Balkan and western Turkey areas, and extending to Calabria in Italy, with yellow (sometimes reddish-yellow), almost sessile round fruits that ripen much later than those of the wild cherry plum. (ii) P. brigantina Vill., a bush (ca. 2 m. high) with similar yellow, late-ripening fruits restricted to a small mountainous area near Briançon, south-east France. Both wild plums are quite similar morphologically. They might represent only geographical races of a single species. Domesticated hexaploid P. domestica had been thought to be a polyploid product of a cross between diploid P. cerasifera [= P. divaricata] plums and tetraploid sloe, P. spinosa L. (Crane and Lawrence 1952, p. 237; Watkins 1995). The latter is a wild, thorny bush widely distributed over central and northern Europe and the cooler and wetter areas in western Asia. However, the available cytogenetic evidence in support of such polyploid origin of the hexaploid domesticated domestica plums is far from convincing. Comparative morphology does not confirm this supposition. The domestica plums look strikingly similar to the cerasifera forms. Both groups intergrade and apparently cross with one another. In contrast, P. spinosa, with its small and very astringent fruits, is morphologically distinct and seems to

be isolated reproductively from the domestica-cerasifera plums. Hybrids between these two groups are almost fully sterile. The hypothesis that hexaploid P. domestica has a CC SS S1S1 genomic constitution, (i.e. contains two genomes (S,S1) contributed by P. spinosa), does not comply with the above facts. As argued by Zohary (1992), it is unlikely that the sloe contributed two genomes (or even a single genome) to the domesticated hexaploid domestica plums. In conclusion, the 6X P. domestica plums seem to be closely related to the 2X, 4X, and 6X P. cerasifera. Together they form what seems to be a P. cerasiferaP. domestica polyploid crop complex. If wild forms of 6X P. domestica existed in south-east Europe and/ or south-west Asia prior to domestication (and they probably did), they should be regarded as the ancestral stock for the development of this fruit crop. However, if as some botanists believe, domestica plums evolved only under domestication, the plausible principal progenitor is the P. cerasifera aggregate. In the Balkans and west Turkey, crosses with 2X P. cocomilia could have enriched variation under domestication. Also the 4X sloe, P. spinosa, might have contributed to the domestic gene pool, not by allopolyploidy and donation of entire genomes, but through secondary hybridization—by rare formation of 5X domestica X spinosa spontaneous hybrids and subsequent introgression.

Archaeological evidence Plum stones appear in several Neolithic and Bronze Age sites in Europe, in countries such as Italy, Switzerland, Austria, and Germany. They are variable in shape, yet they fall within the morphological range of present-day cerasifera and insititia plums (Bertsch and Bertsch 1949; Körber-Grohne 1996). In addition, a large quantity of stones reported as P. domestica subsp. insititia was uncovered in Bronze Age Tell Mudrets (Gudzhova Mogila), Bulgaria (Popova 1995). All these finds seem to represent fruit collected from the wild. In contrast with the paucity of plum remains in Neolithic and Bronze Age contexts, domestica-type stones were repeatedly uncovered in numerous Roman sites in Germany and neighbouring countries (Körber-Grohne 1996; Stika 1996; Willerding 1996). While no archaeobotanical finds preserved remains in 79 BC Pompeii,

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Fig. 38 Plum, Prunus domestica, Roman Period Masada, Israel (photograph kindly provided by M. Kislev, Bar Ilan University).

wall painting there depicts blue, purple, and yellow plums. Feemster and Meyer (2002) identified the blue and purple plums as P. domestica, while the yellow one remains unidentified. They continue to be present also in later periods (Fig. 38). Such appearance suggests cultivation. In summation, we still know very little about the origin of cultivated P. domestica and the beginnings of plum domestication, but since its culture depends on grafting, this fruit tree was probably taken into cultivation together with apples and pears. The earliest literary records of plum planting and grafting are from Roman times.

Cherries: Prunus avium and P. cerasus Cherries are characteristic fruit trees of the cooler, temperate parts of the Old World (Iezzoni et al. 1991; Watkins 1995). Two species of cherries are widely grown: (i) the diploid (2n = 2x = 16 chromosomes)

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sweet cherry Prunus avium L., and (ii) the tetraploid (2n = 4x = 32 chromosomes) sour or morello cherry, P. cerasus L. The sweet cherry, P. avium L., is a rather tall tree (up to 2 m high) with sweet, round, red-black berries. Both the wild forms and the cultivars are largely self-incompatible and require cross-pollination in order to set fruits. Maintenance is by grafting. According to Pliny, this fruit crop was brought to Rome from the Pontus area (southeastern part of the Black Sea basin) by Locullus in 73 BC upon his return from the East (White 1970). The domesticated clones of the sweet cherry are closely related to a group of wild and feral forms of P. avium, which are widely distributed over temperate Europe, northern Turkey, Caucasia, and Transcaucasia (Meusel et al. 1965, Vol. 2, p. 227; Webb 1968; Browicz 1982, Vol. 1, Map 72). The ripe fruits of these spontaneous cherries are smaller than those of the domesticated varieties and usually reach only 10 mm in diameter. They show the same black-red colour of P. avium and have either a sweet or a bitter taste, but are never acid. Also the stones of the wild forms closely resemble those of the cultivars but are usually smaller (7–9 mm in the wild as compared to 9–13 mm in most of the domesticated forms). Both the cultivated and the wild P. avium are diploid (2n = 4x = 32 chromosomes) and largely self-incompatible. The domesticated sour cherry, P. cerasus [= Cerasus vulgaris Miller] is a smaller tree, rarely exceeding 8 m in height, with bright red berries and a characteristic acid taste (Hanelt 2001, vol 1. pp. 505–508). Most cultivars are self-compatible. The crop is closely related to, and fully inter-fertile with, the ground cherry, P. fruticosa Pallas., a wild tetraploid (2n = 4x = 32 chromosomes) self-pollinated shrub or small tree, native to parts of eastern Europe and adjacent south-west Siberia. Both the wild and cultivated are tetraploid, fully inter-fertile, largely self-compatible and known to hybridize with one another (Hanelt 1997). P. fruticosa is likely the direct wild progenitor of the domesticated sour cherry, or at least a major contributor to its formation through crosses with P. avium. Its small fruits have a cherry-like taste but are too astringent to be palatable.

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Archaeological Evidence Cherries were collected from the wild long before their cultivation. The numerous finds of Neolithic and Bronze Age stones in central European settlements (Bertsch and Bertsch 1949) seem to represent wild forms. Cultivated cherries make their appearance rather late. The earliest reliable report of cherry cultivation comes from classical times. Pliny tells that Lucullus, in the first century BC introduced to Rome a superior cherry variety which he had obtained in the Pontus region of the Black Sea (White 1970). Large quantities of cherry stones were found in both Roman and medieval contexts in many European countries (Baas 1980; SchultzeMotel 1968–1994; Kroll 1995–2000).

Latecomers: apricot, peach, and quince Three other members of the rose family, Rosaceae, namely (i) apricot, Armeniaca vulgaris Lam. [= Prunus armeniaca L.], (ii) peach, Persica vulgaris Miller [= Amygdalus persica L., Prunus persica (L.) Barsch.] and (iii) quince Cydonia oblonga Miller [= C. vulgaris Pers.] are not Mediterranean elements. They seem to have arrived to the temperate parts of Europe and the Mediterranean basin from the east, and very probably already as domesticated horticultural crops. Wild forms of apricot, peach, and quince, fully inter-fertile with their respective cultivars, grow in central and eastern Asia. Smaller stands of wild apricot also extend to the Transcaucasia (including Armenia and Dagestan). The wild quince is much more restricted in its geographic distribution. Wild forms of quince thrive in the belt of the Hyrcanian vegetation in the southern part of the Caspian Sea basin. Probably, they arrived in south-west Asia and in Europe via Persia and/or Armenia as crops at a late date. Reliable archaeobotanical and literary evidence on their arrival appears only in classic period (fifth century BC). Apricot Armeniaca vulgaris Lam. (= Prunus armeniaca L.) is a diploid (2n = 2x = 16 chromosomes) fruit-tree, grown for its large, fleshy fruits. Some cultivars are self-compatible, and a few others are self-incompatible. Wild-growing populations of apricot, are fully inter-fertile with the crop. Wild

forms of A. vulgaris, with their relatively small leaves and fruits, are widely distributed over central Asia, (particularly in Tien Shan mountain system and several other mountain ranges, and in north-eastern China, Korea, eastern Siberia, and Ussuria). There, an additional two or three species of Armeniaca occur (Mehlenbacher et al. 1990; Watkins 1995). Smaller stands of wild apricot extend to Armenia (Gabrielian and Zohary 2004) and to Dagestan. Indeed, apricot stones were reported from several ca. 6,000–4,750 cal. BP Eneolithic settlements in Ukraine (Januševič 1978; Pashkevich 2005). In the wild forms, and in many cultivars, the large seeds (enclosed in hard shells) are bitter. Similar to almonds, this bitterness protects the seeds from predation. In some other cultivars this wild-type adaptation broke down. The seeds are palatable and frequently used as nuts. The apricot was apparently grown in China some 3,000 years ago. It is widely assumed that it was introduced into domestication in northern China or eastern Asia (Bailey and Hough 1975; Watkins 1986; Buttner 2001). However, the evidence supporting a Chinese origin is still scanty. We still know very little about the early history of apricot domestication in central Asia, or Armenia—territories where wild forms apricot occur as well. The apricot reached the Mediterranean region relatively late. Very likely, it was introduced into south-west Asia from Iran or Armenia around the third century BP (Hopf 1973b). A few hundred years later apricot cultivation was well-established in Syria, Turkey, Greece, and Italy. Peach, Persica vulgaris Miller [= Amygdalus persica L.; Prunus persica (L.) Barsch.] is a diploid (2n = 2x = 16 chromosomes) fruit crop. Its wild forms occur in the mountainous areas of Tibet and western China. There are records pointing to peach domestication in China as early as 4,000 years old (Hesse 1975; Huang et al. 2008). On the basis of available classical literature, Hedrick (1919, p. 463) concluded that the peach reached Greece from Persia in classical times (at about 300–400 BC). However, a discovery of peach remains in ca. seventh century BC Heraion, Samos (Kučan 1995) indicates that its arrival might have been several hundreds of years earlier. The Romans did not

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cultivate it until AD first century (Huang et al. 2008). Soon after, the peach was established as a valued fruit crop in the Mediterranean basin. Bakels and Jacomet (2003) surveyed the archaeobotanical finds of ‘luxury food’ in Roman Period Europe. They report peach remains in thirty-two sites, mostly in the south; the few peaches in the north are all from military settlements—apparently imported. Roman Period finds are reported also from Israel and Egypt. Quince Cydonia oblonga Miller [= C. vulgaris Pers.] is a large shrub or a small tree, largely confined to the Hyrcanian vegetation belt in the southern coastal areas of the Caspian Sea (Browicz 1996). Quince is a diploid (2n = 2x = 34 chromosomes) monotypic genus (only a single species in its genus) in this restricted territory. Wild forms of C. oblonga are frequent in Azerbaijan, especially in Talish and in Dagestan districts, as well as along the Caspian Sea coastal belt in northern Iran. Wild growing quince populations are reported also from the Kopet Dagh range in South Turkmenia, but they might be feral. Apparently, domestication took place within this restricted area. The relatively large and gritty fruits (poms) of domestic quince clones are widely used in south-west Asia and in the Mediterranean basin. Usually, the fruits are not eaten raw, but cooked to prepare jams, compotes and beverages. So far, there is no archaeobotanical documentation on this fruit crop. However, literary sources indicate that the quince reached the Mediterranean region in classical times. When used as a stock for grafting (for apples, pears, etc.) quince frequently produce a dwarfing effect in the grafted scions, resulting in smaller trees which make it easier to harvest the fruits. Some archaeological find were retrieved mainly in medieval contexts in Germany and other central European countries (Schultze-Motel 1968–1994; and Kroll 1995–2000).

Carob: Ceratonia siliqua The carob, Ceratonia siliqua L. (Caesalpiniaceae), is a characteristic constituent of the evergreen, maquistype vegetation in low-elevation areas in the Mediterranean basin. All over these territories, this evergreen dioecious tree has been extensively culti-

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vated for its pods, which are used both for human consumption and as a sugar-rich animal feed (Batlle and Tous 1997; Zohary 2002). Under domestication, clones bearing fleshier, sweeter pods have been selected. In superior cultivars, the pod’s sugar content can reach 50% or more. In plantations, the majority of the trees are female. This requires the addition of male pollen donors to the plantation in attempt for satisfactory production of pods. Hermaphrodite mutants that do not need crosspollination for fruit set occurred as well, and were protected by growers. With the advent of mechanized agriculture and the availability of relatively low-priced sugar, carob cultivation in the Mediterranean basin shrank considerably. However, the endosperm of C. siliqua seed contains a unique, highly viscose gum (galactomannan carob gum), which is increasingly used as a stabilizer in commercial food products. This new development might cause a comeback of carob cultivation. Similar to most other fruit crops, the domestication of C. siliqua is based on a shift from sexual reproduction (in the wild) to vegetative propagation (under domestication). The carob does not lend itself to simple methods of vegetative propagation. Therefore, the growing of domestic carob clones depends entirely on scion grafting. For this reason, it is likely that this fruit tree was taken into domestication only after the arrival of this propagation technology into the Mediterranean basin. Domestication probably, happened only in the later part of the third millennium BP. The available archaeobotanical documentation is too scanty to suggest when and where this fruit tree could have been taken into domestication (Zohary 2002). Carob pollen grains were detected in a pollen core in the Hula Valley, north Israel (WeinsteinEvron 1983), in sediments belonging to the Early Würm period (ca. 40,000 BP). C. siliqua pollen grains were found also in ca. 14,000 BP Hayonim Cave (West Galilee) in contexts dated ca. 14,000 BP (LeroiGourhan 1981). Two charred carob seeds were uncovered in Pre-Pottery Neolithic contexts at Nahal Oren (Noy et al. 1973), and pieces of charred wood were discovered in ca. 10,000-9,300 BP PrePottery Neolithic A and in ca. 9,300-8,500 BP PrePottery Neolithic B levels in Jericho (Western 1971). Finally, both seeds and wood charcoal of carob were

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uncovered at ca. 8,000–7,500 BP late Pre-Pottery Neolithic Atlit-Yam (Galili et al. 1993). All these finds antedate the beginnings of horticulture in south-west Asia and show that C. siliqua was part of Levantine flora at least since early Paleolithic times. Carob remains continue to appear later, in Pottery Neolithic, Chalcolithic, and Bronze Age contexts (Liphschitz 1987; Kislev 1988; Zohary 2002; Liphschitz, 2008). A few charred seeds were found in Chalcolithic Abior Cave near Jericho (Kislev 1992), Early Bronze Age II Arad (Hopf 1978b) and from Iron Age Ashkelon (Weiss and Kislev 2004). Yet most of its remains (recovered mostly in Israel) come from Roman times or later. Jewish and Roman literary sources provide clearer clues. The tree is not even mentioned in the Old Testament. In contrast, carob cultivation is soundly documented in later Jewish sources—the Mishna (first century BC to AD second century) and the Talmud (third to the fifth centuries AD). During the Mishna time, carob cultivars were extensively grown in Israel and their pods were commonly used as human food (Feliks 1994). The Mishna specifies that the maintenance of these clones depend on grafting. Furthermore, a clear distinction is made between: (i) the superior pods produced by grafted cultivars, which are fit for human consumption, and (ii) the inferior pods borne by non-grafted carobs, which are suited only for animal feeding. In Rome, two contemporary writers, Columella (4 BC to AD 65) and Pliny the elder (AD 23–79), describe the carob and its cultivation at some length. In addition, numerous charred carob pods were uncovered in AD 79 Herculaneum and Pompeii (Meyer 1980). The situation in the western part of the Mediterranean basin is less clear. Wild growing C. siliqua trees abound here. Yet some researchers (see Batlle and Tous 1997) suspect that in the westMediterranean basin the carob is naturalized.

Citrus fruits Domesticated citrus fruits (the genus Citrus L., Rutaceae), had their origin in south-east Asia and in the Indian subcontinent (Davies and Albrigo 1994; Roose et al. 1995). In these parts of the world one still finds the wild relatives from which they could have been derived. Citrus cultigens comprise some

eight main types (citron, lime, lemon, pummelo, bitter orange, orange, mandarin, and grapefruit). All are largely inter-fertile, and when crossed produce a wide range of recombinant and intermediate progeny. Citrus specialists usually refer to each of these main types as a distinct domestic species. However, the majority of them are probably only hybrid products that originated, only under domestication, from much fewer wild stocks; probably from wild forms of citron, mandarin, and pummelo (Scora 1988). Numerous cultivars of citrus are apomictic. They set seed containing asexual, nucellar embryos, which retain the genetic constitution of the mother plant. Such a mode of reproduction is exceptional among fruit crops and it allows the maintenance of desired genotypes by the simple planting of seeds. Apomixis was apparently of major advantage in the early stages of domestication of citrus fruits. By this means, vegetative propagation was possible before the introduction of grafting in the Mediterranean area. The citron Citrus medica L. is the only citrus crop that was grown in south-west Asia and the Mediterranean basin in Greek times. It apparently arrived from India (via Persia), and its fruits, with their characteristic thick aromatic rind, were appreciated medicinally. Jewish literary sources (and archaeological depictions in mosaics and coins) indicate that in Hellenistic times, the citron acquired a religious symbolic role (especially in Sukkot, the Feast of Tabernacles). Theophrastus’ detailed description of its cultivation and propagation leaves little doubt that by the end of the fourth century BC, C. medica was already well established in the east Mediterranean basin. A single archaeobotanical find suggests an even earlier arrival: Hjelmqvist (1979a) uncovered a few charred citrus seeds in ca. 3,200 BP Bronze Age Hala Sultan Tekke, Cyprus. But these remains have not yet been directly dated (by AMS radiocarbon method) to confirm their antiquity. Better identified and dated finds came from Roman sites in Egypt, Mons Claudianus, where entire fruits including seeds were found (van der Veen 2001a), Mons Porphyrites (van der Veen and Tabinor 2007), and Quseir al-Qadim (van der Veen 2003). Several other citrus crops: lemon, C. limon (L.) Osbeck; Seville or bitter orange C. aurantium L.; lime

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C. aurantiifolia (Chistm. & Panz) Swingle; and pummelo C. maxima (Burm.) Merrill., arrived in the Mediterranean basin, apparently in early Islamic times (Amar 2000; Ramón-Laca 2003). Ramón Laca (2003) studied literary sources and concluded that these crops entered Europe via the Iberian Peninsula and Sicily. As for grapefruit (C. paradisi Macfad.), mandarin (C. reticulata Blanco), and sweet orange (C. aurantium L.), she concluded that they arrived to the West only during the last 500–200 years as a result of the trade with the British and Portuguese colonies.

Almond: Amygdalus communis The domesticated almond, Amygdalus communis L. (Rosaceae) [= Prunus amygdalus Batsch., P. dulcis (Miller) D.A. Webb], is a widely grown nut tree in the Mediterranean basin (Watkins 1986), and probably one of the earliest, fruit tree domesticants in Old World agriculture. Among the Rosaceous fruit crops, almonds are the earliest to flower (Plate 17). They thrive best in the relatively warm Mediterranean-type climate. Compared with olives and grapes, almonds can endure somewhat drier conditions. Most domesticated almonds, as well as the wild forms, are diploid (2n = 2x = 16 chromosomes), self-incompatible, and cross-fertile. Some cultivars, such as the Apulia or the Bari groups, carry a mutation that renders them self-compatible. Wild almonds are characterized by the amygdaline contents of their seeds, which render them bitter and poisonous. The main trends of evolution under domestication were selection for mutant types with non-poisonous, sweet seed, larger drupes, and softer, thinner shells. Almonds cannot be propagated by rooting, and its modern cultivation depends mainly on grafting. In contrast with most other fruit trees, almonds are also planted from seeds. These seedlings produce considerably varied fruits, in shape and size of the fruits, in hardness and thickness of the shells, and in seed bitterness. Most of the non-bitter individuals are very tasty. The non-bitter condition is governed by a single dominant mutation. Therefore, 75% or more of the progeny raised from seed harvested in non-bitter almond plantations produce sweet fruits (Heppner 1923; Heppner 1926; Spiegel-Roy and

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Kochba 1981). For this purpose, farmers traditionally collected seeds from trees producing sweet seeds. Seed planting necessitates rogueing of unwanted bitter individuals. This has been a traditional way of almond cultivation in numerous localities in the Mediterranean basin and in southwest Asia.

Wild ancestry The genus Amygdalus L. comprises some twenty-six vicarious species distributed over south-west and central Asia, as well as south-east Europe (Browicz and Zohary 1996). The almond is closely related to an aggregate of wild forms native to the Levant countries. These wild almonds, which are placed taxonomically within A. communis complex, fall into two intergrading eco-geographical races: (i) relatively large wild and weedy forms, A. communis subsp. spontanea, thrive in Mediterranean environments (350–800 mm annual rainfall) and resemble closely the domesticated varieties in flower and leaf morphology, in early blooming, and in their climatic requirements; and (ii) more xeric, smaller wild forms, A. communis subsp. microphylla (Post) Browicz & D. Zohary [= A. korschinskyi (Hand.-Mazz.) Bornm.], which occupy drier ‘steppe forests’ or steppe-like environments. These almonds have smaller leaves and fruits compared to their more robust Mediterranean counterparts. Wild forms, placed taxonomically in subsp. spontanea, are identified as the main progenitor of the domesticated varieties (Browicz and Zohary 1996). They differ from the cultivars mainly by their smaller fruits, harder shells with fewer pits and their intensely bitter seed. The bitterness represents a chemical defence. It is brought about by the presence of the glycoside amygdalin, which becomes transformed into deadly prussic acid (hydrogen cyanide) after crushing, chewing, or any other injury to the seed. The consumption of a few dozen bitter nuts yields enough prussic acid to prove fatal for human beings. Domestication of the almond focused upon selection of non-bitter almonds. Sweet

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cultivars, known as var. dulcis, contain only traces of amygdalin. Some of them are homozygous, others are heterozygous for the dominant mutation. Like numerous other fruit trees, A. communis comprises a complex of interconnected wild forms, weedy derivatives, and cultivars in its main area of distribution. In the Levant, wild forms of subsp. spontanea grow in Mediterranean maquis and oak park-forest formations, thriving best on rocky slopes and escarpments facing south. ‘Weedy’ forms often colonize secondary habitats such as neglected plantations, edges of terrace cultivation, and roadsides. The latter are frequently feral derivatives, or products of spontaneous hybridization between cultivated and wild forms. Several other Amygdalus species, growing outside the Levant, are quite close morphologically to the crop and are interconnected to it by spontaneous hybridization (Browicz and Zohary 1996). Most prominent among them are: A. webbii Spach, native to the Aegean basin and south Italy; A. fenzliana (Fritsch) Lipsky in north-east Turkey, adjacent Armenia, and Caucasia, A. kuramica Korsh. in northeast Afghanistan, and A. bucharica Korsh. in Tadzhikistan and Uzbekistan. All these wild almonds were very likely isolated geographically from A. communis in the past. However, the spread of almond cultivation into their territories brought them in contact with the crop. As evident from spontaneous hybrids encountered in places of contacts, this superimposition initiated hybridization. The extent of the gene-flow between the domesticated crop and these wild almonds is hard to assess. Similar to the apple or pear, the presence of intermediates and recombinants indicates that introgression from the local wild species could have facilitated the development of locally adapted A. communis cultivars and/or helped in the establishment of local feral A. communis stands.

Archaeological evidence Almonds were apparently collected from the wild long before the plant’s domestication. It is unclear how to differentiate between wild and domesticated almond shells in archaeological finds (endocarp). Charred wild almond shells were uncovered in Epi-Palaeolithic Ohalo II, Israel (Kislev et al.

1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004, 2008), and in Epi-Palaeolithic Öküzini cave, Turkey (Martinoli and Jacomet 2004), and in Mesolithic and Neolithic levels at the Franchthi Cave, Greece (Hansen 1991a). Charred PPNA almonds were found in Netiv Hagdud, Jordan Valley, Israel (Kislev 1997; Hartmann 2006), and in Jerf al Ahmar on the Euphratus river, Syria (Willcox 2002; Willcox et al. 2008, 2009), and Early PPNB Çayönü, Turkey (van Zeist 1972; van Zeist and de Roller 1991–2, 2003). A rich find is available from ca. 10,000 cal BP Late Natufian Hallan Çemi Tepesi, South Turkey (Rosenberg et al. 1995). Here a large quantity of charred wild fruits was uncovered, suggesting the use of wild almond seeds. (Perhaps latent toxicity was mitigated, either by leaching the water-soluble prussic acid, or by extraction and use of almond oil). In Knossos, Crete, almonds played an important role during the Neolithic Period, from Aceramic Neolithic, ca. 8,650–8,400 cal BP on (Sarpaki 2009). In later periods, almond remains come from late Neolithic Dhali Agridhi, Cyprus (Stewart 1974). Aceramic Neolithic Çatalhöyük, Turkey (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007) Tepe Musiyan, Iran (Helbaek 1969), as well as Late Neolithic Dimini (Kroll 1979), Sesklo (Kroll 1981a), and Sitagroi (Renfrew 1979, Table 3) in northern Greece. They also occur in late Neolithic/ Chalcolithic Hacilar, Turkey (Helbaek 1970). Recently, Martinoli and Jacomet (2004) differentiated between wild Amygdalus and Prunus on the basis of their morphology. Almonds are also reported from Early Bronze Age Bab edh-Dhra, Jordan (McCreery 1979). Here they appear with numerous remains of grapevine and olive, which probably represent domesticated trees. These could have been grown either under irrigation (near to the site) or under rain-dependent conditions (at higher elevations nearby). Remains of almond shells continue to appear in Bronze Age sites in south-west Asia, Egypt, and Greece. They were found in ca. 1,325 BC Tutankhamun tomb in Egypt (Germer, 1989a; Hepper 1990; de Vartavan et al. 2010; see also Fig. 39). From classical times on, A. communis is recorded as a characteristic element of Mediterranean horticulture (White 1970).

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Fig. 39 Desiccated almond shells, Tutankhamun’s tomb, New Kingdom, Egypt (Photo: A. McRobb, with kind permission of F. Nigel Hepper, The Royal Botanic Gardens, Kew, UK).

In conclusion, almonds appear to have been members of the earliest domesticated fruit-tree assemblage in the Old World. They were probably introduced into domestication in the eastern part of the Mediterranean basin, apparently at about the same time that the olive, grapevine, and date palm were domesticated, i.e. in the Chalcolithic Period, in about 6,300–5,300 cal BP. The almond does not lend itself to vegetative propagation from suckers, cuttings, or rooting. However, unlike most Mediterranean fruit trees, attractive almonds can be raised from seeds. This indicates that the almond could have been domesticated even before the introduction of grafting.

Walnut: Juglans regia The common or Persian walnut tree, Juglans regia L. (Juglandaceae), is a traditional nut of Old World agriculture. This large tree produces beautiful hard timber in addition to its fruits. The fruit is a drupe, having a fleshy, green, outer layer that encloses the ‘nut’, which dries and falls off at maturity. The hard shell of the nut (the endocarp) encloses a single large, edible seed, which is rich in oil. Juglans regia is

diploid (2n = 2x = 32 chromosomes) and is a windpollinated, monoecious tree. Juglans regia grows wild in mesic, temperate, deciduous forests of the Balkans, eastern Turkey, Armenia, Azerbaijan, north Iran, the south Caspian region, the Caucasus, and central Asia. It reappears in the Tien Shan province of western China. Wild trees produce variable small fruits (2–3 cm in diameter), which have relatively thick shells. Their seed is as tasty as that of the domesticated tree. The domesticated varieties have larger fruits, which are 3–6 cm in diameter. They are grown today in the native distribution range of J. regia as well as in central and western Europe and in the more xeric environments in the Mediterranean basin and western Asia. In the latter, the walnut thrives best in cool, hilly areas and its cultivation usually benefits from supplementary irrigation in summer. Like the almond, the walnut does not lend itself to rooting by cuttings or to propagation by suckers. Cultivation depends mainly on grafting of selected clones but, like almond, this nut tree can also be raised from seed. However, by this method trees vary considerably in the size and shape of their fruit and in the thickness of the shells.

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Information pertaining to the time and place of walnut domestication is still inadequate. Earliest remains of wild J. regia appear in ca. 10,650–10,250 cal BP Mesolithic Balma Margineda, Spain (Marinval 1995), ca. 9,450–8,450 cal BP Late/Final Neolithic Can Hasan III, Turkey (Hillman 1972; Hillman 1978), and ca. 7,550–6,450 cal BP Early Neolithic Sammardenchia and adjacent sites, northern Italy (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007). Such remains continue to appear in several European and south-west Asian countries in the later Bronze and Iron Ages. As for the Levant, Liphschitz (2008) reports wood remains from ca. 1,550–1,150 cal BC Middle Bronze onward in northern Israel. Eventually, J. regia became more common from Roman time onwards. Bakels and Jacomet (Bakels and Jacomet 2003) surveyed the archaeobotanical finds of ‘luxury food’ in Roman Europe, and report walnut remains in seventy-two sites. Roman period finds reported also from Israel and Egypt. Such sites are: the Cave of the Pool and Abior, the Sandal and the Spear caves, Judean Desert, Israel (Zaitschek 1962; Kislev 1992; Simchoni and Kislev 2009), and Bernike, Quasir al-Qadim, Mons Claudianus, and Mons Porphyrites, Egypt (Cappers 1996, 2006; van der Veen 1999, 2001a; Vermeeren and Cappers 1997/2002a). Greek, Roman, and Jewish literary sources indicate that the walnut was already widely grown in Classical times in the Mediterranean Basin and south-west Asia. J. regia cultivation started probably before Roman times, particularly in southwest Asia. Palynological data (Bottema 1980; Bottema and Woldring 1984) indicated that J. regia disappeared from south-east Europe and south-west Turkey during the last glaciation (Würm glaciation), and reappeared in the Balkans and west Turkey not before the middle of the fourth millennium BP. This late reappearance suggests that J. regia did not return to these areas as a post-glacial wild element but was reintroduced by humans. If this is true, the spontaneously growing walnuts in the Balkans and central Europe represent feral derivatives of cultivated walnuts introduced by humans as recently as the Bronze Age. This point to north-eastern Turkey, the Caucasus, and north Iran as the most plausible area of walnut domestication.

A large quantity of desiccated and charred walnut fruits were found in Roman Masada, Israel. They include both thick (majority) and thin (minority) shells (Kislev pers. comm.). This indicates that the mutant of ‘thin shell’ was already present some 2,000 years ago.

Chestnut: Castanea sativa The sweet chestnut, Castanea sativa L. (Fagaceae), is a large tree and a valued nut crop in the humid parts of northern Turkey and southern Europe. Chestnuts trees have been utilized extensively in various south European countries and in Asia Minor for managed coppicing and pollarding stands, as well as for their nuts. The nuts formed an important part of the diet in traditional farming communities and were also used as animal feed. Chestnut production in southern Europe and south-west Asia suffered debilitating damage from the attacks of two fungal diseases (chestnut blight and ink root disease), introduced from America in the 1890s. As a result of these diseases, chestnut stands in many places were decimated and production fell to 10.2% of the former level. Chestnut domestication is based primarily on the selection of clones producing large and tasty nuts, and their maintenance by grafting. Wild or naturalized stands which thrive on steep slopes and acid soils are frequently grafted in situ with scions producing superior nuts. Castanea L. comprises some 10–12 species spread over the northern hemisphere (Richardson 1986). The domesticated chestnut is closely related to a variable aggregate of wild and feral forms which occur in the northern parts of the Mediterranean basin, north Turkey, and Caucasia (for details, see Meusel et al. 1965, Vol. 2, p. 121; Browicz 1982, Map 48). Such wild forms also penetrate deeply into the climatically milder areas in central and western Europe. Because of their close taxonomic affinity with the domesticated clones, botanists include these spontaneous chestnuts within C. sativa. As with the walnut, our knowledge of the place of origin and time of domestication of C. sativa is still inadequate. Charred remains of this lovely nut tree start to appear in several European countries in the later Iron Age. Remains become more common

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from Roman time onwards (for enumeration of the finds see Schultze-Motel 1968–1994; Kroll 1995– 2000, as well as the online databases http://www. cuminum.de/archaeobotany/database/; http:// www.archaeobotany.de/database.html). The palynological record indicates that C. sativa disappeared (or almost disappeared) from southern Europe during the Würm glaciation, and that its pollen reappeared in quantity in Anatolia and Greece only at about 1,500–1,300 BC (van Zeist 1980). An even more recent increase in pollen frequency was found in Italy and other west Mediterranean sites (van Zeist 1980). This suggests that the comeback of C. sativa to Europe was assisted by humans, and again points to north Turkey and the Caucasus as the most probable area for the initial chestnut domestication. However, as van Zeist (1991) noted, chestnut remains were retrieved from ca. eleventh century BC Greifensee-Bochen near Zürich; and its presence was attested (by charcoal remains) in ca. 1,500 BC Monte Leoni, north Italy. These finds suggest that C. sativa might have survived the last glaciation in some local refugia in Europe, and that its reoccupation might not necessarily have been exclusively from the east.

Like almonds, hazels were also grown from seed obtained from superior individuals. Although sexual reproduction results in a wide variation in nut size and shape, all nuts, even the small ones, are tasty. It is not clear where and when the domestication of C. avellana was started, but apparently clones of this shrub were already planted by the Romans (White 1970, p. 259). Corylus maxima Mill. is fully inter-fertile with and frequently included taxonomically within C. avellana. It is native to the Balkans, north Turkey, and Caucasia. The fruit is somewhat larger than in typical C. avellana. C. maxima has very attractive nuts (Lagerstedt 1975) which were, and still are, extensively collected from the wild, and superior plants were taken into cultivation (as clones). Because of the bigger fruit size and better yields, planting of maxima-type clones spread considerably in recent times and often replaced C. avellana in west and central Europe and Turkey. Today the world production of hazel nut is primarily based on C. maxima cultivars, (which are called the true filberts), or on modern hybrid derivatives obtained from crosses between C. maxima, C. avellana, and several additional Corylus species.

Hazelnut: Corylus avellana

Pistachio: Pistacia vera

Hazelnut (also known as filbert), Corylus avellana L., belong to Corylus L. (Betulaceae)—a genus of about fifteen species of deciduous shrubs and trees. (Wright 1986). This common hazel of Europe and west Asia, appears as a shrub or small tree. Its nuts, measuring 10–25 mm in length, are covered with an involucre. C. avellana is a common component of the broad-leaved oak and beech forest belt of temperate Europe, Caucasia, north Turkey, and the Caspian belt of Iran (Meusel et al. 1965, Vol. 2, p. 118; Browicz 1982, Map 48). The tasty, rounded or ovoid nuts are easily collected and shelled. Its remains have been repeatedly retrieved from many Neolithic, Bronze Age, Classical, and Medieval contexts all over Europe and south-east Asia. The European hazel was also taken into cultivation and planted both for its nuts and for a supply of branches used for the preparation of hurdles and walking sticks. Superior clones were traditionally kept by the layering of branches.

The pistachio nut, Pistacia vera L. (Anacardiaceae), ranks among the most drought resistant of nut trees. Domesticated varieties of this crop are extensively grown in Iran, south-eastern Turkey, central and south-west Asia, for its oval (ca. 1–2 cm long), tasty nuts. The shell encloses a single oil-rich seed and splits longitudinally, along its lateral suture, when the nut is ripe. Pistacia vera is a dioecious, diploid (2n = 2x = 30 chromosomes) wind pollinated nut tree. Fruit-bearing female clones have been traditionally planted, intermixed with male individuals. Domestication depends on grafting. Frequently, the P. vera cultivars are grafted in situ on stocks of other wild Pistacia species (P. palaestina, P. atlantica, P. khinjuk, or P. terebinthus). They may be pollinated by several other indigenous pistachio species. Pistacia vera grows wild in north-east Iran, north Afghanistan, and in the middle Asian republics— Uzbekistan, Tadzhikistan, Kirgizia, and the southern most parts of Turkmenia and Kazakhstan—where

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this xeric tree constitutes a conspicuous element in the local semi-arid, ‘steppe forest’ vegetation belt (Browicz 1988, map 5; Zohary 1996, Fig. 5). Wild forms have smaller (but edible) fruits. They are also collected and consumed by the local people. Recent molecular research revealed the occurrence of genetic diversity and affinity among wild forms and cultivars in the genus Pistacia (e.g. Shanjani et al. 2009; Pazouki et al. 2010). The distribution area of the wild forms of P. vera indicates that this tree could have been brought into cultivation in central Asia. Indeed, the earliest archaeological documentation of its use comes from Bronze Age Djarkutan, Uzbekistan (Miller 1999), and from late Neolithic and Bronze Age Tepe Yahya, Iran (Costantini and Costantini-Biasini 1985).

Whether these finds represent collection from the wild or domestication is hard to say. But the total dependence of pistachio on grafting today suggests relatively late domestication (the earliest literary source of grafting technology is Theofrastus fourth century AD). P. vera was introduced into the Mediterranean region under the reign of Emperor Tiberius (Buttner 2001, p. 1057). There are no clear signs of P. vera cultivation in south-west Asia or in southern Europe before classical times, although a single P. vera half-shell was reported by Renfrew (1973) in late Neolithic Sesklo, Greece. Another single shell was found by Kroll (1983) in Bronze Age Kastanas, Greece. In our view both these finds are questionable. They probably represent more modern intrusions into the reported layers.

C H A PTER 7

Vegetables and tubers

Compared to cereal grains and stones of fruits, vegetables and tuber crops are highly perishable. In archaeological contexts, their soft parts are only rarely charred and preserved. It is therefore not surprising that very few remains of vegetables and tubers have been found in archaeological excavations. The only exceptions in south-west Asia are Egypt, and to some extent caves in the Judean Desert and the Dead Sea Rift Valley, where numerous desiccated vegetable and tuber remains survived because of the extreme dryness. The available archaeobotanical data on the early phases of vegetable domestication in the Levant are therefore fragmentary. However, this is partly compensated for by evidence from Mesopotamian Bronze Age literary sources, and by drawings and descriptions found in Egyptian tombs. The combined evidence shows that, since the start of the fourth millennium BP, vegetable gardens constituted an integral element of food production both in Mesopotamia and in the Nile Valley. Furthermore, most of the vegetables grown at these early times have been satisfactorily identified: watermelon, melon, leek, garlic, onion, lettuce, and chufa appear to be major constituents of Bronze Age vegetable production. The number of vegetable crops grown in humanmade gardens in South-west Asia, Egypt, and Europe, seem to have increased considerably in the third millennium BP (= first millennium BC). An early written source is the list of plants grown in the garden of Merodach-Baladan in Babylonia (ca. 2,720 BP). The list enumerates some of the earlier vegetables (lettuce and garlic), as well as beet, turnip, and cress (Körber-Grohne 1987). Towards the end of the third millennium BP the number of vegetable crops

seems to have increased considerably. Greek, Roman, and Jewish classic writers were familiar with a large number of vegetables (for reviews see Lenz 1859; White 1970; Körber-Grohne 1987; Small 2006). At that time cabbage, turnip, beet, celery, carrot, parsnip, asparagus, and several other herbs, had already become part of the vegetable garden. Significantly, almost all these new cultigens have had closely related wild forms growing in the same geographical regions. Therefore, it seems they could have been taken into cultivation in that part of the world. Ethnographic data shows (see, for example, Ertuğ, 2004, 2009) that wild-growing vegetables take part in the diet of traditional farming communities, even at the present time. There is no doubt that this tradition of collecting wild vegetables predates agriculture. Recently, seeds of several Mediterranean herbs and vegetables (such as celery, dill, and parsnip) have been found in Neolithic layers in Europe (see below and in Chapter 8). It is uncertain whether these are wild forms or domesticated ones. They might have been local wild-growing plants, garden planting, or imported from the Mediterranean region.

Watermelon: Citrullus lanatus The watermelon, Citrullus lanatus (Thunb.) Matsum. & Nakai [= C. vulgaris Schrad.], is belongs to the gourd family (Cucurbitaceae), having crawling liana. It is an annual summer crop, widely seed planted in warm regions, and greatly appreciated for its large sweet fruit with a useful source of water (90% or more). It is diploid (2n = 2x = 22 chromosomes) and largely cross-pollinated. In addition to the pulp, the oil-rich seeds are also consumed.

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The crop is closely related to four diploid wild taxa, placed in the genus Citrullus (Jeffrey 1975: Singh 1990): (i) C. colocynthis (L.) Schrad., or colocynth, a perennial wild watermelon bearing relatively small (5–8 cm), bitter fruits, which is widely distributed over the deserts and semi-deserts of Africa, the Mediterranean basin, and south-west Asia; (ii) C. ecirrhosus Cogn., a similar perennial watermelon, restricted to the Kalahari Desert; (iii) C. lanatus, wild growing, annual watermelon forms, with somewhat larger fruits, which also thrive in south-west Africa; and (iv) C. rehmii De Winter, an additional annual watermelon species reported from Namibia (de Winter 1990). All these wild types are inter-fertile with the crop (Shimotsuma 1963; Navot and Zamir 1987; de Winter 1990). Yet C. lanatus and C. rehmii stand somewhat closer to the cultigens in morphology and molecular composition. On the basis of this evidence, some botanists concluded that the lanatus-type annual forms constitute the wild stock from which the crop evolved, and that south-west Africa probably was the place where watermelon domestication took place. Currently, the available archaeological information does not seem to support south-west Africa as the locale of watermelon domestication. Soundly identified remains of domestic watermelon appear in Egypt already at the start of the fourth millennium BP. As far as we know, at this time farming was not yet practiced in south-west Africa (see chapter 9). In other words, when watermelon was already a crop in Egypt, the cultural environment for its domestication did not yet exist in the area where we find its closest wild relatives today. This contradiction is hard to explain. The intensely bitter fruits of the colocynth have long been valued for their strong purgative effects. The fruits are still collected by nomads in the Sahara and south-west Asia and sold to pharmacies. After special preparation, the seeds are sometimes used as human food (Hedrick 1919; Osborn 1968). Crosses between wild C. colocynthis and the domesticated watermelon have shown that the shift from bitter fruits to non-bitter ones is caused by a recessive mutation in a single gene (Navot et al. 1990).

Archaeological evidence Significantly, the characteristically small seed of C. colocynthis appear in several early Egyptian, Lybian, and south-west Asian sites, including Neolithic Armant (Lityńska-Zając 1993), ca. 5,650–5,850 BP Nagada (Wetterstrom 1993, 1998), and ca. 5,500– 5,300 BP Hierakonpolis (El Hadidi et al. 1996) in Egypt, Pre-Pottery Neolithic B Nahal Hemar Cave, Israel (Kislev 1988), and a series of sites ranging from ca. 5,800 BP to Roman times in Libya (van der Veen et al. 1996). These finds indicate that humans probably used the wild colocynth prior to its domestication. Watermelon was grown in the Nile Valley at least since the start of the fourth millennium BP. Several finds of its characteristically large seeds (5–8 mm in diameter), as well as a single find of leaves, are reported by Keimer (1924, pp. 17–18), the oldest of them dating from the twelfth dynasty (ca. 3,985–3,773 BP). Numerous desiccated watermelon seeds were found in Tutankhamun tomb, ca. 1,325 BC (Germer 1989b; Hepper 1990; de Vartavan et al. 2010). The Agricultural Museum of Dokki, Cairo, has a sample of watermelon seed retrieved from the New Kingdom Thebes (Darby et al. 1977). Van Zeist (1983) confirmed these early finds and reported on the presence of watermelon remains in the foundation deposits of the walls of two eighteenth-dynasty (ca. 3,500 BP) temples near Semna, Nubia. Wasylikova and van der Veen (2004) describe watermelon remains as a funeral gift (together with domestic grapes, olives, figs, dates, and rushnuts) in the same temples. Finally, Cox and van der Veen (2008) reported a significant size increase of watermelon seeds from the Roman to the Islamic period at Quseir al-Qadim, Egypt. They also suggest that the site dwellers had eaten the seeds during the Islamic period, but not during the Roman period.

Melon: Cucumis melo The domesticated melon, Cucumis melo L., is a second cucurbit crop, which was probably brought into cultivation in south-west Asia or in Egypt at a relatively early date. It is a very variable crop that includes both: (i) sweet fruited varieties (melons or muskmelons), and (ii) non-sweet green-fruited

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forms (chate-melons, orfakus in Arabic). The latter are rare today, frequently bear bent fruits, and are consumed like cucumbers. They were traditionally referred to as C. chate Hasselqu., or C. melo L. var chate (Hasselqu.) Naud. The wild ancestry of the domesticated crop (2n = 2x = 14 chromosomes) is well established (Jeffrey 1980; Sebastian et al. 2010). The domesticated varieties show close morphological resemblance and full inter-fertility with a variable group of wild and weedy annual and perennial melons distributed over the subtropical and tropical parts of Asia, Africa, and Australia (Kirkbride 1993). The more xeric type (frequently referred to as C. callosus (Röttl.) Cong.), with ovaries covered with dense long hairs, is native to central and south-west Asia. It is closest to the melon cultivars raised in western Asia and the Mediterranean basin. The Indian and east Asiatic cultigens show closer affinities to a second, more tropical, wild melon race in which the young ovaries are covered by a short, close pubescence.

Archaeological evidence There are few archaeological remains of melons. However, they seem to indicate that C. melo was cultivated in Egypt in the Bronze Age. Melon seeds were discovered in Predynastic, ca. 5,650– 5,450 cal BP Hierakonpolis (Fahmy 1995, 2003, 2005; El Hadidi et al. 1996; Fahmy et al. 2008), and ca. 5,450–5,300 cal BP Maadi (van Zeist and de Roller 1993, 2003). Illustrations of offerings, as well as faience models of what are clearly the bent fruits of the green-fruited melons (var. chate) decorate several ancient Egyptian tombs from the Old Kingdom period on (Keimer 1924). Three carbonized seeds are available from Late Bronze Age Tiryns, Greece (Kroll 1982) and a few others from ca. 4,000 BP Shahr-i Sokhta, east Iran (Costantini 1977). The melon, C. melo was the principal (and the oldest) cucurbitaceous vegetable of ancient Mesopotamia, at least from the beginning of the fourth millennium BP on (van Zeist and WaterbolkRooijen 2003, p.74). The ‘small’ and ‘finger’ varieties mentioned by Stol (1987b) were probably cucumber-like, non-sweet forms. The ‘large’ and

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the ‘ripe’ forms could represent more advanced non-sweet or even sweet melons. The sweet melons are the melopepo of the Greeks. Indeed, numerous C. melo seeds were uncovered in twenty-seventhth century BP Heraion, Samos (Kučan 1995). A third Old World principal cucurbit, namely the cucumber, Cucumis sativus L. (diploid, 2n = 2x = 24 chromosomes), does not belong to the indigenous vegetable ensemble of south-east Asia and the Mediterranean basin. Wild forms of this vegetable, namely C. sativum L. var. hardwickii (Royle) Alef. occur in the Himalayas and in adjacent territories east of this mountain belt (Jeffrey 1980; Kirkbride 1993; Sebastian et al. 2010). These wild forms were reported also from the province of western Ghats, Peninsular India (Kuriachan and Beevy 1992). Most likely, C. sativus was taken into cultivation in India and arrived in the Mediterranean basin rather late. Exactly when this happened is still hard to say, especially since the seeds of C. sativus and C. melo are quite similar in their external features. They do differ, however, in their seed coat anatomy (Kučan 1995). In the future, this might help to trace the arrival of the cucumber into south-west Asia, although typically cucumbers are eaten unripe before the seeds mature. Recently, Janick, Paris, and Parrish (2007) reviewed the occurrence of cucurbits in Classical and Jewish literature, as well as in artistic evidence from around the Mediterranean basin. They described the use and growing of melon, bottle gourd (Lagenaria siceraria, used mainly as vessels or utensils), and watermelon as the most important cucurbits in this area during the Roman Period. However they found no convincing evidence for the existence of C. sativus around the Mediterranean in this time period, and they conclude that the cucumis of Columella and Pliny was not actually a cucumber, but C. melo subsp. melo Flexuosus Group (snake/chate melons, or fakus). It seems, therefore, that the cucumber arrived to the area only after the classical times.

Leek: Allium porrum The garden leeks (kurrats in Arabic), placed in Allium porrum L. (Liliaceae), ranked as one of the popular vegetables in the old world. They are mostly tetraploid (2n = 4x = 32 chromosomes) and

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seed planted. Ancient Egyptian wall carvings and drawings, as well as several finds of dried specimens (Täckholm and Drar 1954; Keimer 1984) show that A. porrum was part of the Egyptian food production from the fourth millennium BP onwards. Outside Egypt, there are few remains. These include wild or domesticated material from Early and Middle Bronze Age Jericho (Hopf 1983). Available texts indicated that leek was grown in Mesopotamia at the beginning of the fourth millennium BP (Stol 1987a). Leek is certainly a Mediterranean or south-west Asian element and its wild progenitor is well identified (McCollum 1976). The more robust varieties grown for their thick ‘pseudostem’ (A. porrum L. senu stricto) and the slender leafy forms (sometimes referred to as A. kurrat Schweinf.), are all closely related to, and inter-fertile with, the wild and weedy tetraploid forms of wild Allium ampeloprasum L., which is widely distributed in the Mediterranean basin.

millennium BP) and later tombs (Täckholm and Drar 1954; Germer 1989b). Outside of Egypt, there are few ancient garlic remains. A large number of carbonized garlic cloves were uncovered from the fourth millennium BP Tell ed-Der, Iraq (van Zeist and Vynckler 1984). Also, garlic is well recognized linguistically in numerous cuneiform records that indicate garlic was cultivated in Mesopotamia since at least the beginning of the fourth millennium BP (Stol 1987a). In the last decade, garlic finds were reported from some European sites as well. The wild ancestry of domesticated garlic has not been definitely established so far. The common sterility of the cultivars makes it difficult to identify of its wild progenitor. Because of this feature, the garlic crop has not yet been fully cross-tested with its wild relatives. The cultivars bear the closest mor-

Garlic: Allium sativum Garlic, Allium sativum L. (Liliaceae), was also an early constituent of the south-west Asian vegetable garden. Garlic is a diploid (2n = 2x = 16 chromosomes) bulb crop (and see Hirschegger et al. 2010 for molecular and taxonomic treatment). In contrast to the leek, garlic is maintained almost totally by vegetative propagation (planting of individual cloves). Up until now, the prevailing view was that most garlic cultivars are totally sterile, and their flowers do not set seeds. However, a few recent collections obtained in central Asia and the Caucasus (Kamenetsky 2007) was found to be almost seedfertile or semi-fertile. The earliest, excellently preserved, desiccated garlic remains were discovered in the ‘Cave of the Treasure’ near Ein Gedi, Israel, dated the ca. 5,970 BP Middle Chalcolithic period (Zaitschek 1961; M. Broshi pers. comm.). Egypt provides early archaeological evidence for this crop as well. Similarly excellently preserved garlic (Fig. 40) was found in ca. 1,325 BC Tutankhamun’s tomb (Germer 1989a; Hepper 1990; de Vartavan et al. 2010). Several wellpreserved dry remains of garlic are available from other Egyptian eighteenth dynasty (mid-fourth

Fig. 40 Desiccated bulbs of garlic, Tutankhamun’s tomb, New Kingdom, Egypt (Photo: H. Barton, with kind permission of Griffith Institute, Ashmolean Museum, Oxford).

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phological, as well as molecular resemblance to Allium longicuspis Regel, known from several scattered locations in east Turkey, Iran, and central Asia. This taxon has been regarded (Stearn 1978) as the most probable candidate for the ancestry of domesticated garlic clones. However, similar to the cultivars, A. longicuspis inflorescence is bulbiferous and mostly sterile. As argued by Mathew (1996) this raises serious doubts whether it could be regarded as a wild ancestor of domesticated A. sativum. Alternatively, it might represent cultivated garlic planted (and abandoned) by nomads. For this reason, other fully fertile close relatives, namely A. tuncelianum (Kollmann) N. Özhatay, B. Mathew & Siraneci (native to south-east Turkey), and A. macrochaetum Boiss. & Hauskn. (native to southwest Asia) were suggested by Mathew (1996) as additional candidates for the ancestry of domesticated garlic. Also, A. truncatum (Feinbr.) Kollman & D. Zohary (native to the Levant) is morphologically close enough to be suspected. All these three wild Alliums smell like garlic.

cal records of onions are not yet available. Cuneiform sources indicate (Stol 1987a) that this vegetable has been grown in Mesopotamia since the fourth millennium BP. Like garlic, wild onion is apparently not a Mediterranean element (Brewster, 1994). Its progenitor has not been convincingly identified. The crop shows close morphological and molecular affinities with two central Asiatic wild onions: A. oschaninii O. Fedtsch., which grows in north Afghanistan, Tadzhikistan, and neighbouring Uzbekistan, and A. vavilovii M. Pop. & Vved., native to the Kopetdag mountains, Turkmenia (Hanelt 1990). Crosses between A. oschaninii and domesticated A. cepa gave rise to sterile F1 hybrids (Hanelt 1985; Hanelt et al. 1992), indicating that this wild onion is not a direct source from which the vegetable has been derived. In contrast, A. vavilovii is reported to be almost fully inter-fertile with the crop (Raamsdonk et al. 1992). There are doubts whether these vavilovii collections represent genuine wild material or only feral derivatives of the crop.

Onion: Allium cepa

Lettuce: Lactuca sativa

Onion, A. cepa L., is the third member of its genus that was apparently highly appreciated as a garden crop in south-west Asia and the Mediterranean basin. It is a diploid (2n = 2x = 16 chromosomes), cross-pollinated, seed-planted bulb crop. Wall carvings and drawings depicting offerings of onions and illustrations showing how onions are planted and watered appear in numerous Egyptian tombs from the Old Kingdom onwards (Germer 1989b; Hepper 1990). These include the Unas (ca. 4,420 BP) and Pepi II (ca. 4,200 BP) pyramids. Well-preserved onion remains are available from the eighteenthdynasty tombs in Egypt (Täckholm and Drar 1954; Germer 1989b). They are complemented by somewhat later finds, including bulbs placed in mummies (and see detailed survey in Murray 2000b). Egypt continues to be the main locale for onions in later periods, like Roman Iron Age Berenike and Shenshef (Vermeeren and Cappers 2002; Cappers, 2006), Kellis (Thanheiser et al. 2002), Mons Claudianus (van der Veen 2001a), and Quseir alQadim (van der Veen 2003; van der Veen et al., 2009). Outside of Egypt, undisputed early archaeobotani-

Lettuce, Lactuca sativa L., from the Compositae (Asteraceae) family, is an annual, diploid (2n = 2x = 18 chromosomes), self- pollinated vegetable crop. It is definitely a west-Asiatic and Mediterranean element. The main use of lettuce is for salads. Its wild ancestor is fully identified (Zohary 1991; de Vries 1997; Koopman et al. 1998). The crop shows close morphological resemblance to a series of six to seven wild lactuca species native to south-west Asia. Its closest wild relative is weedy L. serriola L., which abounds in the Mediterranean basin and south-west Asia. It is an aggressive summer weed in much of the subtropical and temperate climate zones of the world. Wall illustrations of rosettes of a tall, large vegetable with subulate leaves appear in numerous Old Kingdom and Middle Kingdom tombs and monuments in Egypt (Keimer 1924; Körber-Grohne 1987, plates 81–82). Most archaeobotanists agree that these leafy vegetables depict the garden lettuce. Unfortunately, the available illustrations are not yet complemented (in Egypt) by dry remains of lettuce leaves or lettuce rosettes, although seeds of

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lettuce have been reported (Germer 1985). Outside Egypt, indisputable records of L. sativa cultivation appear only in Greek and Roman literature (Pliny, for example, mentions eleven distinct forms).

Chufa or rush nut: Cyperus esculentus Chufa or rush nut, Cyperus esculentus L. (Cyperaceae), ranks among the oldest domesticated plants in Egypt. It is a vegetatively propagated sedge crop. Its small (1.5–2.5 cm) ellipsoid-globose tubers appear in large quantities in Egyptian archaeological contexts from the pre-dynastic Amratian period (ca. 5,900–5,650 BP) onward (Täckholm and Drar 1950; Wetterstrom 1993, 1998). Dry remains of Cyperus esculentus tubers were discovered as foundation deposits in two temples near Semna, Nubia, ca. 3,500 BP. They indicate that Chufa was already a crop by the fourth millennium BP (van Zeist 1983). In addition, chufa tubers are frequently illustrated in tombs (Darby et al. 1977). Cyperus esculentus represents a local domestication. The wild and weedy forms of this sedge (var. aureus Richt.) with its smaller and more fibrous tubers, grow in the Nile Valley, in wet places, and is collected or grown by the local people. While Chufa was an important food element in ancient Egypt throughout dynastic times, it did not spread much beyond this area. There are almost no contemporary records of this tuber-bearing sedge from other parts of the Old World.

Cabbage: Brassica oleracea European cabbage, Brassica oleracea L., from the Cruciferae (Brassicaceae) family, was a wellestablished Mediterranean garden vegetable and oil crop already during Greek and Roman times (Körber-Grohne 1987). This extraordinarily polymorphic crop includes (U 1935; Hodgkin 1995) leafy kales, hearting cabbages, broccoli, cauliflower, Brussels sprouts, kohlrabi, and marrowstem kale (the latter is an animal feed). In classical times leafy, cauline forms prevailed. Theophrastus (372–287 BC) mentions three kinds of cabbage: curly leaved, smooth leaved, and a wild-type. Pliny the elder (AD 23–79) reported on a wider variety of domesticated forms, which might also have included hearting

types and broccoli-like forms. Types like cauliflower and Brussels sprouts appeared much later. The B. oleracea cultivars are closely related to, and fully (or almost fully) inter-fertile with a varied aggregate of perennial, herbaceous, largely selfincompatible, diploid (2n = 2x = 18 chromosomes; genomic constitution CC), wild cabbage races. These wild varieties (the B. oleracea ‘cytodeme’) grow mainly on winter-rain coastal cliffs and similar maritime habitats along the Atlantic coast of Europe, the Mediterranean basin, and the Canary Islands (Snogerup et al. 1990; Gustafsson and Lannér-Herrera 1997). The following closely related ten wild oleracea taxa have been recognized in this complex; many of them are narrow endemics: (i) Wild B. oleracea subsp. oleracea, native to the Atlantic coast of Spain, France, UK, and Helgoland; (ii) B. montana Pourret [= B. oleracea subsp. robertiana (Gay) Bonnier & Layens] native to the coast of north-east Spain and the French and the Italian Rivieras; (iii) B. rupestris Rafin. and (iv) B. villosa Biv. (sensu lato), both in west Sicily; (v) B. incana Ten. in east Sicily, the west coast of Italy, and several Adriatic islands; (vi) B. macrocarpa Guss., endemic to Isole Egadi near Sicily; (vii) B. insularis Moris in Corsica, Sardinia, and Tunisia; (viii) B. cretica Lam. in Greece, the Aegean islands, Crete, and south-west Turkey; (ix) B. hilarionis Post in Cyprus; and (x) B. bourgeaui (Webb) Kuntze in the Canary islands. As to the location of B. oleracea initial domestication, two possible ancestral populations were suggested, but no conclusive evidence can be offered currently. Based on morphological similarities, cytogenetic affinities, and molecular comparisons, the Atlantic B. oleracea subsp. oleracea seems to be the closest wild relative of the domesticated varieties of cabbage (Song et al. 1990; Hodgkin 1995; Gustafsson and Lannér-Herrera 1997). The other wild vicarious taxa are more distantly related. Yet some of them (e.g. B. cretica) could also have been involved in the evolution of the crop. On the other hand, (Maggioni et al. 2010) gathered linguistic, literary, and historical information, and found out that they indicate the domestication of B. oleracea in the ancient Greek-speaking area of the central and east Mediterranean. The molecular survey of Christensen et al. (2011) characterizes diversity and asseses the genetic structure among seventeen

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accessions of kale landraces, cultivars, and wild populations from Europe. The results of this research, however, were inconclusive regarding the ability to indicate a single geographical location for kale’s centre of origin.

Turnip: Brassica rapa Turnip, Brassica rapa L. [= B. campestris L.], of the cabbage family (Cruciferae/Brassicaceae), is a cross-pollinated vegetable widely cultivated in China and south-west Asia for its swollen hypocotyle (root turnip), its leaves, and for its oil (turnip rape or oil-seed rape, subsp. rapa). Other groups of turnip cultivars, such as Chinese cabbage and Peking cabbage (subsp. pekinensis) have evolved in east Asia. The wild progenitor of this crop complex is well identified. Wild forms, and particularly weedy races of B. rapa are distributed all over the Mediterranean basin, temperate Europe, Siberia, and in south-west and east Asia. They are fully inter-fertile, diploid (2n = 2x = 2 chromosomes), and chromosomally identical with the cultivars. Because of the vast distribution of the wild turnip forms, and the differences between the western and eastern cultivars, it seems possible that B. rapa was domesticated independently both in the belt of Mediterranean agriculture and in China. (Note that the Chinese finds are not treated in this book).

Archaeological evidence Similar to many other Old World vegetables, turnip is poorly represented in archaeological contexts. It is, however, soundly identified linguistically. Thus, written sources partly compensate for this deficiency. Seeds of B. rapa were found in several Neolithic and Bronze Age lake-dwelling sites in Switzerland and neighbouring countries. They indicate that at these early times, wild turnip was an element of the weed flora of arable lands in this part of Europe (Körber-Grohne 1987; Jacomet et al. 1991; Brombacher 1995; Jacomet 2007). Probably, the seeds of these weedy forms were collected for their oil, but B. rapa does not seem to have been a crop at that time. In contrast, root turnip, grown for its swollen hypocotyle, was part of the Old World vegetable garden. The earliest sign of its cultivation

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is found in the list of plants grown in the garden of King Merodach Baladan II (722–711 BC) in Babylonia (Körber-Grohne 1987). From Hellenistic times onwards, turnip is treated by several Greek and Roman authors who wrote about agriculture (for review see Lenz 1859). It is also referred to in the Jewish Mishna (Feliks 1983). From Roman Iron Age onward, B. rapa is present in few archaeological sites across Europe. Swede, Brassica napus L., is a root and oil-seed cruciferous crop similar looking and closely related to the turnip. Yet swede is a tetraploid cultigen (2n = 4x = 38 chromosomes). It probably originated under domestication, by hybridization (and chromosome doubling) between the diploid cabbage B. oleracea and the diploid turnip B. rapa. This root occurs in archaeological finds came from Roman Period onward.

Beet: Beta vulgaris Beet, Beta vulgaris L. (Chenopodiaceae) is a variable, multi-purpose crop, comprising four principal groups of cultivars (Körber-Grohn, 1987; Frese 2003). The oldest cultigens are the leaf beet (with edible leaves) and the garden beetroot (with swollen hypocotyle). Both are used as food. Their domestication was followed by the development in Europe of large-rooted fodder beet and, at the start of the nineteenth century, by breeding of sugar beet. The crop is closely related to and fully interfertile with a group of chromosomally homologous, diploid (2n = 2x = 18 chromosomes), wild taxa, placed in section Beta of the genus Beta (Ford-Lloyd and Williams 1975; Ford-Lloyd 1995). Closest to the cultigens is a variable aggregate of wild and weedy beets regarded by these authors as wild races of the crop complex, namely: (i) B. vulgaris subsp. maritima (L.) Arcangeli [= B. maritima L.] which is widely distributed in the Mediterranean basin, south-west Asia, and the Atlantic coastal belt of Europe; (ii) B. vulgaris subsp. macrocarpa (Guss.) Thell. [= B. macropcarpa Guss.], native to the warmer, more arid parts in the Mediterranean basin; and (iii) B. vulgaris subsp. adanensis (Pamukç.) Ford-Lloyds & Williams [= B. adaninsis Pamukç.], which thrives in the Mediterranean parts of Turkey. These wild beets constitute the general stock from which the domestic

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beet could have been derived. Subsp. maritima is most probably the principal progenitor. Beet is poorly represented in archaeological contexts, and only a few of its remains have been uncovered (Knörzer 1991; Kreuz 1994–5; Kučan 1995). Moreover, charred fruit remains of domestic forms are very similar to those of wild forms. However, beet is linguistically well identified. Early written sources mentioning this vegetable compensate, to some extent, for the paucity of the archaeological evidence. The earliest written documentation on beet cultivation comes from eighth century BC Babylonia (Körber-Grohne 1987). Theophrastus (ca. 372–287 BC) describes this vegetable as having a thick taproot similar to that of the radish. Roman and Jewish literary sources indicate that in the first century BC domestic beet was represented in the Mediterranean basin by leafy forms (chards, mangold) and probably also by beetroot cultivars. Beet remains are available also from a few sites in Egypt: a desiccated part of a flowering stem was found in the thirrd-dynasty Saqqara pyramid at Memphis (Lauer et al. 1950; Germer 1985). Four charred beet fruits were retrieved from Neolithic Aartswoud, north Netherlands (Pals 1984). Since wild forms of B. vulgaris abound in these places, it is hard to say whether the finds represent domestic forms or wild material. The majority of the scanty remains of charred beet fruits come from Roman sites in Germany (Knörzer 1991; Stika 1996). Significantly, at least some of these German sites lie outside the distribution area of wild beets, and suggests they were domesticated. The appearance of domesticated beet in Roman Germany corroborates the contemporary literary sources.

Carrot: Daucus carota The carrot, Daucus carota L. (Umbelliferae/Apiaceae), is another vegetable which was grown in the Mediterranean basin already in classical times. There are both annual and biannual forms of this crop. Two main types of domesticated carrots are recognized: (i) ‘eastern domestic forms’ with dark purple, anthocyanin-coloured roots, and (ii) ‘western forms’ with orange, carotene-coloured roots. Members of the first group, grown mainly in

Afghanistan, Iran, and Turkey, often have branched roots and are considered the more primitive cultivars. Carotene-coloured cultivars characterize carrot cultivation in the Mediterranean basin and temperate Europe. All commercial cultivars belong to this group. The available archaeological information on carrot is deplorably fragmentary. Consequently, the tracing of its history under domestication relies, to a large extent, on written sources. The earliest satisfactory evidence on carrot domestication comes from Greek and Roman writers (Körber-Grohne 1987). The plant named by Dioscorides (60 AD) as ‘staphylinos’ is apparently D. carota. The telltale dark spot in the centre of the inflorescence, mentioned by Dioscorides identifies it as such. We are told that the staphylinos’ aromatic root is being used both as food and for medicine, and that garden-grown plants are tastier than the wild ones, but less potent medicinally. The illustration in Dioscorides codex, drawn in Constantinople ca. AD 500, shows a carrot plant with a thick, orange-coloured root, indicating that carotene-containing cultivars already existed at that time. Carrot mericarps became more abundant in Middle Ages archaeological contexts throughout Europe. The domesticated carrot is closely related to, and fully inter-fertile with, a variable assemblage of wild and weedy carrot races, all included by taxonomists in the D. carota species complex (Heywood 1983). All are diploid (2n = 2x = 18 chromosomes) and largely cross-pollinated herbs. About a dozen eco-geographic wild subspecies have been recognized in D. carota, most of them in the Mediterranean basin and south-western Asia. Weedy forms have an even wider distribution and extend over most of temperate Europe. They often grow close (or even within) carrot cultivation. Under such conditions spontaneous hybridization between the domesticated carrot and its weedy relatives is quite frequently.

Celery: Apium graveolens Celery, Apium graveolens L. (Umbelliferae/Apiaceae), is presently grown for its fleshy petioles (var. dulce) or for its swollen hypocotyle (var. rapaceum).

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However, less advanced cultivars grown for their leaves and/or thick aromatic hypocotyle have had a much longer history under domestication. They were part of the Mediterranean vegetable garden assemblage in classical times, appreciated both as medicinal and garnishing herbs. The wild ancestor of celery is well identified (2n = 2x = 22 chromosomes). Wild forms of A. graveolens, fully interfertile with the cultivars, thrive in marshy places all over the Mediterranean basin and southwestern Asia. They extend north, along saline coastal marshes, as far as the British Isles and Denmark. Celery leaves and inflorescences were part of the garlands placed in ca. 1,325 BC Tutankhamun tomb, and with the mummy of Kent (second dynasty, ca. 1,000 BC) in Egypt (Germer 1985; Hepper 1990; de Vartavan et al. 2010), and celery mericarps were discovered in seventh century BC Heraion, Samos (Kučan 1995). However, since A. graveolens grows wild in these areas it is hard to decide whether these remains represent wild or cultivated forms. The information available from Switzerland is even more puzzling. Here, celery mericarps were found in several late Neolithic lake-shore settlements (e.g. Jacomet 1988, 2004; Jacomet et al. 1989; Brombacher 1997; Favre 2002; Brombacher et al. 2005); i.e. places in which A. graveolens does not grow wild today. This suggests transport and planting by humans. Yet it is hard to believe that this plant was under domestication in such early times. Convincing evidence on celery domestication is available only from classical times (Körber-Grohne 1987). By Roman times A. graveolens was part of the vegetable garden in the Mediterranean basin. Moreover, at that time its use seems to have expanded north of the Alps. Celery were retrieved from several Roman sites in Italy, Switzerland, and Germany (Stika 1996; Willerding 1996). Evidence of celery also arrives from ca. 2,700–2,400 cal BP Zinchecra, Libya (van der Veen, 1992a, 1992b).

Parsnip: Pastinaca sativa Parsnip Pastinaca sativa L. is another umbelliferous (Umbelliferae/Apiaceae) root vegetable grown in the Mediterranean basin and in temperate Europe since classical times (Körber-Grohne 1987; Riggs

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1995). Its white, thick taproot has been used as a cooked vegetable and also for animal fodder. Parsnip is a biennial (2n = 2x = 22 chromosomes) crop, grown from seed. It is a European and west Asian element. Wild and weedy forms of this vegetable (bearing much thinner and less succulent roots) are widely distributed over temperate Europe and north Turkey. They are also found in some of the relatively moister parts of the Mediterranean basin itself. In some areas (especially in the north) wild-looking P. sativa might be feral. Remains of parsnip have been retrieved from Roman sites in Germany and several other European countries (Körber-Grohne 1987; Kreuz 1994–5; Stika 1996). Altogether the archaeological evidence on parsnip is limited. Similar to several other vegetables, Greek and Roman literary sources constitute a major source of information on its early use. There are some difficulties in distinguishing between parsnip and carrot in classical writings since both vegetables seem to have been sometimes called ‘pastinaca’, yet each vegetable appears to be well under domestication in Roman times.

Asparagus: Asparagus officinalis Asparagus officinalis L. (Liliaceae) is a rhizomatous, perennial vegetable cultivated for its tender, young shoots (‘spears’). The crop is dioecious, and diploid (2n = 2x = 2 chromosomes). It is a Mediterranean and European element. Wild forms of A. officinalis are widely distributed, particularly in slightly moist habitats in the Mediterranean basin, in adjacent parts of south-western Asia, and extend to numerous parts of temperate Europe. Cytogenetically, the variable wild forms of A. officinalis have not been studied extensively, but they are known to contain both diploid (2n = 2x = 2 chromosomes) and tetraploid (2n = 4x = 40 chromosomes) forms. The crop is most likely derived from a diploid wild stock. Asparagus is a relatively large genus containing some sixty species. Only A. officinalis is cultivated as a vegetable, while several other members of this genus are used as ornamentals. Traditionally, the young shoots of wild A. officinalis (as well as

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those of several other Mediterranean Asparagus species) have been collected from the wild (Ertuğ 2009). This practice survives today in Turkey, and some other countries. Greek and Roman literary sources indicate (Körber-Grohne 1987) that by 200 BC, A. officinalis was grown as a garden vegetable

in Greece. It was appreciated for its medicinal qualities and for its delicious taste. Evidence on its use in Greece is soon followed by substantial written information on its cultivation in Rome. Evidence of older asparagus cultivation is almost nonexistent.

C H A PTER 8

Condiments

Like the vegetables, several condiments indigenous to west Asia and the Mediterranean basin entered domestication rather early. The tradition of their planting and usage is well documented in classical Greek, Roman, and Jewish sources. Only few remains of this group of crops have been discovered in archaeological contexts. Many condiments were used not only for their seeds but also for their green leaves, which like vegetables rarely survived archaeologically. Because of the rarity of condiment finds, our knowledge of the beginning of their domestication is insufficient. Many of the available records come from Egypt, where the combination between the use of a wide range of plants and desiccation in desert environment, leads to the survival of a wide range of material (see Murray 2000a, b, c, as an excellent review of the Egyptian evidence). Frequent finds of several condiments and herbs of Mediterranean origin, e.g. parsley, dill, celery, and lemon balm (Melissa officinalis), start to appear in lake-shore settlements in Switzerland already in Late Neolithic, ca. 6,000 cal. BP. It is unknown yet whether these species were locally grown or were imported from regions closest to the Mediterranean coast (Jacomet 2007). Interestingly, the remains of four condiments (coriander, cumin, dill, and black cumin) were discovered in several Hellenistic and Roman sites in Egypt, like the remote desert quarry of Mons Claudianus, Egypt (Van der Veen 1998, 2001b), the Red Sea harbour Berenike (Vermeeren and Cappers 2002; Cappers 2006), and Quseir al-Qadim (Van der Veen 2003, 2009). These finds indicate that during the Roman Period they were considered essential for Romans’ diet. As Livarda and van der Veen (2008) noted, the availability of condiments

increased significantly with the Roman Empire, and to some extent characterizes this period. The following sections review the available evidence on what seem to have been the first condiments of the Old World.

Coriander: Coriandrum sativum Coriander, Coriandrum sativum L. (Umbelliferae/ Apiaceae), is an annual, largely diploid (2n = 2x = 22 chromosomes), self-pollinated crop, grown for its aromatic seeds and foliage (Diederichsen 1996; Small 1997). It is one of the earliest condiment crops of the Old World (Gabrielian and Zohary 2004), and it was known in classical times. Plant remains and linguistic evidence indicate that its use started much earlier. Today C. sativum occurs spontaneously over wide areas of Old World agriculture and it is hard to define exactly where this plant is wild and where it only recently established itself as a weed or as a naturalized element. In south-west Asia and Armenia, C. sativum seems to grow wild in rocky openings, in oak park-forests, oak scrub, and steppelike formations. Probably, these territories were the source of both the cultivated forms and the weedy races.

Archaeological evidence Fifteen desiccated coriander mericarps were retrieved from Pre-Pottery Neolithic B Nahal Hemar Cave (Kislev 1988), and eleven such mericarps from ca. 8,000–7,500 BP PPNC Atlit-Yam (Kislev et al. 2004), Israel. If they are not intrusive, they represent the earliest archaeological find of this condiment. Early

163

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finds of coriander came also from several Late Neolithic sites: Tell Hammam et-Turkman, Syria (van Zeist 2003), Kapitan Dimitrievo, Bulgaria (Marinova 2006), and Poduri, Rumania (Cârciumaru and Monah, 1985; Monah and Monah 2008). In the latter, the ca. 6,650–6,350 cal. BP Eneolithic stratum, coriander fruits were found mixed with those of Sambucus nigra, inside a painted amphora and wooded cask. A large sample (about half a litre, the main item in several baskets) of round, small, mericarps of C. sativum was found in ca. 1,325 BC Tutankhamun’s tomb, Egypt (Renfrew 1973; Germer 1989a; Hepper 1990; de Vartavan et al. 2010). Because coriander does not grow wild in Egypt, its presence in this tomb, as well as in several other Egyptian sites, suggest that this condiment was under cultivation at that time. Coriander remains were also discovered in the fourth millennium BP Tell ed-Der, Syria (van Zeist and Vynckler 1984), and in several Bronze Age sites: Sitagroi (Renfrew 1973) and Akrotiri, Thera (Sarpaki 1992a), Greece, Ulu Burun shipwreck, Turkey (Haldane 1993; Ward 2003), and Umm el-Mara, Syria (Schwartz et al. 2000). A single half-fruit of C. sativum was found in Late Bronze Age Apliki, Cyprus (Renfrew 1973). Coriander was also known from ca. 1,200–500 B.C. Iron Age Deir Alla, Jordan (Neef 1989), and late Assyrian Nimrud (Helbaek 1966b). It was also found in several Iron Age and Roman sites in Egypt: Berenike and Shenshef (Vermeeren and Cappers 2002), Kellis (Thanheiser et al. 2002), Mons Claudianus (Van der Veen 2001b), and Quseir al-Qadim (Van der Veen 2003, 2009).

Cumin and dill: Cuminum cyminum and Anethum graveolens Two additional aromatic umbellifers (Umbelliferae/ Apiaceae)—cumin, Cuminum cyminum L., and dill, Anethum graveolens L.—were part of Greek and Roman agriculture. Wild forms of cumin are unknown in south-west Asia, but occur in central Asia. Wild and weedy types of dill are widespread in the Mediterranean basin and in West Asia. Earliest cumin seeds were uncovered in ca. 8,000– 7,500 BP PPNC Atlit-Yam, Israel (Kislev et al. 2004), and later in the fourth millennium BP Tell ed-Der, Syria (van Zeist and Vynckler 1984). Later, it was

found as well in Iron Age Deir Alla, Jordan (Neef 1989). In Egypt, a few seeds of cumin were found in New Kingdom Deir el Medineh (Germer 1985), and a basket full of cumin was included in the burial of Kha, architect of Amenophis III, eighteenth dynasty (Wilson 1988). Cumin was found also in several later Iron Age and Roman Age sites in Egypt: Berenike and Shenshef (Vermeeren and Cappers 2002; Cappers 2006), Kellis (Thanheiser et al. 2002), and Mons Claudianus (Van der Veen 2001b), Egypt. Earliest remains of dill was found in several late Neolithic lake-shore settlements in Switzerland (Jacomet and Brombacher 2005; Jacomet 2007). Several twigs of dill were found in the tomb of Amenophis II, eighteenth dynasty, ca 1,550–1,292 BC (Germer 1985) and this condiment is also reported from seventh century BC Heraion, Samos (Kučan 1995). Later it was found in ca 700–400 BC Early Garamantian Zinchecra, Libya (van der Veen 1992a), in Roman Iron Age Berenike and Shenshef (Vermeeren and Cappers 2002; Cappers 2006), and Iron Age Mons Claudianus (Van der Veen 2001b), Egypt. This evidence suggests that already in the fourth millennium BP, both cumin and dill were under cultivation. However, it is difficult to conclude where and when domestication of this condiment began.

Black cumin: Nigella sativa Black cumin, Nigella sativa L., of the buttercup family Ranunculaceae, has been another traditional condiment of the Old World during classical times (Small 1997). Its black seeds were extensively used to flavour food. Archaeological information on the early cultivation of black cumin is scanty, but seeds of N. sativa were reported from ancient Egypt, including ca. 1,325 BC Tutankhamun tomb (Germer 1989a; Hepper 1990; de Vartavan et al. 2010), and Iron Age Mons Claudianus (Van der Veen 2001b). They also appear in Iron Age Deir Alla, Jordan (Neef 1989). Linguistic considerations suggest that this plant was also grown in ancient Mesopotamia (Thompson 1949). Small (1997) identified the use of Black Cumin in Sanskrit, in India.

CONDIMENTS

Wild and weedy forms of N. sativa grow in grain and fallow fields and occur in south Turkey, Syria, north Iraq, as well as in several adjacent territories. This indicates that south-west Asia could have been the place of domestication of this condiment.

Saffron: Crocus sativus Saffron, Crocus sativus L., of the Iris family Iridaceae, is another member of the early group of condiments. The long, scarlet lobes harvested from its flowers were highly valued for flavouring foods and for colouring them golden-yellow. Saffron was extensively grown in south-west Asia and the Mediterranean basin in classical times and it maintained its role as a very, expensive condiment until the beginning of the second century. Recently, production became increasingly impractical because the collection of the styles requires a vast amount of labour. Today saffron cultivation is declining and Crocus survives only as a relic

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crop, despite its position as one of the most expensive spices (Winterhalter and Straubinger 2000). No remains of C. sativus have been traced in archaeological excavations and indeed the chances for the preservation of the delicate stigmas seem fairly low. However, Minoan (ca 1645 BC) pottery and frescoes (e.g. the renowned ‘The Gathering of the Crocus’ from Akrotiri, Santorini) depict Crocus flowers with long exserted, red, stigmatic branches (Mathew 1977; Negbi 1999), and these probably represent C. sativus. Also linguistic evidence suggests that saffron was brought into cultivation before classical times. The cultivated saffron is an autumn-flowering, sterile triploid (3n = 3x = 24 chromosomes) maintained by vegetative propagation. It is morphologically closest to the Greek species C. cartwrightianus Herbert, and is possibly a clonal selection from this wild diploid (2n = 2x = 16) species (Mathew 1977, 1999; Rashed-Mohassel 2006).

C H A PTER 9

Dye crops

A group of dye plants, native to south-western Asia and the Mediterranean basin had probably entered domestication already before classical times. They are amply documented in Greek, Roman, and Jewish literary sources. Woad (Isatis tinctoria L.), dyer’s rocket (Reseda luteola L.), Madder (Rubia tinctorum L.), Safflower (Carthamus tinctorius L.), and indigo (Indigofera tinctoria L.), were characteristic dye crops of Old World agriculture, and their dye stuffs were extensively used to colour textiles and leather. They continued to be highly appreciated crops until the end of the nineteenth century. Then, the invention of low-priced synthetic dyes caused their collapse. In a matter of a few decades, the traditional dye crops of the Old World became commercially redundant and their cultivation died out. Today they represent agricultural drop-outs. Most of the cultivars are extinct. Only a few forms survive in remote farming communities, in botanical gardens, or in the hands of hobbyists. To-date, only few remnants of dye crops have been uncovered in archaeological excavations. This rarity apparently stems from the use of perishable vegetative parts which are rarely preserved, rather than seeds and fruits, for dye production. Consequently, we know very little about the beginnings of dye-plant domestication. Early literary sources, and some analyses of dye elements colouring ancient Egyptian textiles (Eastwood 1984; Germer 1992) make it clear that in the fourth millennium BP, dye crops were used extensively to colour textiles and leather (Krochmal and Krochmal 1974). Some twisted and dyed flax fibers were recently reported in Upper Palaeolithic, ca. 30,000 year-old, Dzudzuana Cave, Georgia (Kvavadze et al. 2009), but raised severe criticism of their validity (Bergfjord et al. 2010; Kvavadze et al. 2010). 166

Woad: Isatis tinctoria Woad, a member of the mustard family (Cruciferae/ Brassicaceae), was a principal dye crop in Europe and in south-west Asia. Until the end of the nineteenth century, woad was cultivated extensively (by seed planting) for its blue indigo dye, which was widely used to colour textiles. The pigment was extracted from the leaves of the vegetative parts of the plants. To obtain the dye, the leaves were dried, powdered, and fermented in alkaline medium—a tedious and very smelly process (Guarino et al. 2000). Today, natural indigo is a commercial rarity. In the textile industry it was mostly replaced by synthetic aniline dyes. Woad—Isatis tinctoria L.—is usually a biennial, cross pollinating, tetraploid (2n = 4x = 28 chromosomes, Darlington and Wyle 1955) plant, with characteristic pale yellow flowers and winged fruits. It is native to south-west Asia and the Aegean area, but it also extends to temperate Europe. Some of the present weedy populations of I. tinctoria (particularly those of temperate Europe) may be feral derivatives of the former crop. Recent research on the genetic variation and population structure of Eurasian collections revealed that the origin of the domesticated crop might have been western and central Asia (Spataro et al. 2007). Similar to madder (see below), classical literary sources show that in the third millennium BP, woad was extensively used in south-west Asia, in the Mediterranean basin, and in temperate Europe (Körber-Grohne 1987). In Neolithic Çatalhöyük (Fairbairn et al. 2002), few seeds of cf. Isatis sp. were recovered, although their identification and use is uncertain. Zech-Matterne and Leconte (2010)

DYE CROPS

compiled recent archaeobotanical and other evidence from north-eastern Europe and raised the possibility of local cultivation of woad in the fifth to the fourth century BC in northern Gaul. In addition, indigo dye (indigotin) was detected in several bluecoloured ancient Egyptian textiles found in eighteenth dynasty (1,370 BC) Tell el Amarna (Eastwood 1984; Germer 1992). While it is impossible to distinguish chemically between indigo extracted from Isatis or from Indigofera (p. 168), only the first seems to have been present in the Mediterranean basin in classical times. Indigofera arrived later.

Dyer’s rocket: Reseda luteola Dyer’s rocket or weld Reseda luteola L. (Resedaceae) is the source of a very intense flavon-type yellow pigment extracted from its roots, stems, foliage, and flowers. This plant was extensively cultivated (by seed planting) until the beginning of the twentieth century when the natural dye was replaced by cheaper synthetic yellow dyes. Dyer’s rocket is an annual or a biennial plant with erect stems and with yellow inflorescences. The plant is apparently indigenous to the east Mediterranean basin and to south-west Asia and became naturalized far beyond its original native geographic range. Historical sources report that R. luteola was harvested in the middle of its flowering season, dried, and sold to textile dyers for dye extraction. It is one of the better natural sources of yellow colour, valuable because it can be used with woad to produce green, an uncommon colour in dyes (Shewry et al. 1997). Classical literary sources show that dyer’s rocket was domesticated and extensively used already in the Old World in the third millennium BP (Körber-Grohne 1987). Like woad and madder, it was probably taken into cultivation even earlier. From Late Bronze Age on, it was found in various sites in west Europe: Belgium, Netherlands, Germany, France, and UK (see Schultze-Motel 1968–1994; Kroll 1995–2000, as well as the online databases http://www.cuminum.de/archaeobotany/database/; http://www.archaeobotany.de/ database.html

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Madder: Rubia tinctorum Madder, Rubia tinctorum L. (Rubiaceae) was, until the end of the nineteenth century, one of the most appreciated dye plants of south-west Asia and Europe. It was extensively grown (by vegetative propagation) all over this area for its rhizomes from which a brilliant red pigment (alizarin) was extracted. Alizarin was widely used to colour linen, wool, cotton, and leather. The dye becomes fixed to the textile fibres only after their treatment with a mordant (alum salts). Furthermore, mordanting with different metals produces different hues (aluminium alum induces dark red coloration; iron alum results in brown-red colour; chromium alum produces a red-violet colour). Pliny (Natural History XXXV xlii) described the technique of madder dyeing in Egypt eloquently: ‘They employ a very remarkable process of colouring textiles. After rubbing the cloth, which is white at first, they saturate it not with the dye—but with mordants that are calculated to absorb the colour. This done, the textiles still unchanged in their appearance are plunged into a cauldron of boiling dye and are removed the next minute fully coloured. It is a remarkable fact, too, that although the cauldron contains a uniform dye, the material taken out is of various colours—according to the nature of the mordants that have been respectively added to the cloth. These colours will never wash out.’

Madder is a perennial herb with characteristic whorls of lanceolate leaves and climbing or staggering scabrous stems that can reach the length of 1–2 m. Its wild forms are native to south-west and central Asia (F. Ehrendorfer, pers. comm.). Spontaneous populations of R. tinctorum thrive also in the Mediterranean basin, mostly in hedgerows, waste ground, and margins of cultivated fields. Possibly, they represent feral derivatives of former cultivation. It is still impossible to conclude where and when madder was taken into domestication or when the alizarin colouring technology was developed. Chemical analysis detected madder dye in redcoloured flax textiles retrieved from eighteenth dynasty (1,370 BC) Tell el Amarna, Egypt (Eastwood 1984; Germer 1992). Various Greek, Roman, and Jewish literary sources show that at the end of the third millennium BP, R. tinctorum was extensively

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cultivated in Persia, Anatolia, and the Mediterranean basin (Körber-Grohne 1987). Evidently, domestication of madder started long before these times. Synthetic alizarin was invented in Germany in 1869. In a matter of only a few decades, the low-cost synthetic equivalent replaced the much more expensive natural dyestuff and caused a collapse of madder cultivation.

True indigo: Indigofera tinctoria In India, as well as in several other tropical and subtropical parts of south Asia and Africa south of the Sahara, the traditional source for indigo blue dye was not woad but several members of the genus Indigofera, from the Papilionaceae/Fabaceae family of the Leguminosae. Prior to the invention of synthetic indigo, a common cultigen was I. tinctoria L. Wild forms of this dye crop occur in India, and this plant was probably domesticated (propagated from seeds) in this subcontinent (Lemmens and WesselRiemens 1991). Several other species of Indigofera, among them I. arrecta Hochst., native to eastern and southern Africa, were also extensively cultivated. In the Indian subcontinent, the pigment extracted from the leaves of Indigofera crops was commercially so valuable, that it was referred to as the ‘king of dye-stuffs’. Sound documentation on its cultivation is available only after the Arab conquest of Egypt.

Safflower: Carthamus tinctorius Safflower, Carthamus tinctorius L. (Compositae/ Asteraceae), was a traditional dye plant of the Old World. Its yellow-red flowers were used extensively to dye textiles and to colour foods. Carthamus is basically an Irano-Turanian genus comprising thirteen to fourteen annual species of thistles (Hanelt 1963). With the recent development of synthetic dyes, the importance of this crop declined considerably. Since the 1950s, safflower acquired a new use and has emerged as a modern industrial oil crop (Knowles and Ashri 1995). The crop is taxonomically closely related to a group of 3–4 diploid (2n = 2x = 24 chromosomes) wild species, placed in section Carthamus, native to south-west and central Asia (Hanelt 1963; Knowles

and Ashri 1995). C. tinctorius was cross-tested with two wild and weedy members of this section, namely: (i) C. persicus Willd.[=C. flavescens Willd.] that occurs in the arid and semi-arid regions of the Levant countries, the Syrian desert, south Anatolia and upper Mesopotamia; and (ii) C. oxyacanthus M. Bieb., that grows in the Trans-Caucasus, Iran, Afghanistan, and Central Asia. Both species were found to be inter-fertile and chromosomally homologous with the crop. C. tinctorius is probably also inter-fertile with a third diploid wild member of this section, namely C. gypsicola Iljin, a geographically much more restricted species, which grows on gypsum-rich substrates in the southern Trans-Caucasus, the southern Caspian basin, and in Middle Asia. In contrast, members of other taxonomic sections in Carthamus are far more distant, and hard or impossible to cross with the domesticated safflower. These three taxa, therefore, can be the progenitor stock of the domesticated plant. Chapman et al. (2010, 2007) conducted recent DNA sequence studies in an attempt to trace the variation and the origin of domesticated safflower. They suggest that C. tinctorius is most likely derived from wild C. palaestinus. However, C. palaestinus is apparently a synonym for C. persicus (Greuter 2006–2009; Flann 2009). Marinova and Riehl (2009) reviewed recently the rich find of safflower crop in archaeological assemblages. Carthamus sp. occurs in the region in several Neolithic and Chalcolithic sites. C. tinctorius achenes appear first in ca. 5,000 cal. BP Early Bronze Age sites in Syria, from where it spread later to Turkey, Bulgaria, and Serbia, to Egypt, the Aegean, and south-eastern Europe. They suggest that the use of safflower for oil production started almost from the beginning of its cultivation. In Egypt, seeds of safflower were found in Late Bronze Age, ca. 1,325 BC Tutankhamun tomb (Germer 1989a; Hepper 1990; de Vartavan et al. 2010). Well-preserved garlands composed of C. tinctorius flowers were found adorning eighteenthdynasty (middle fourth millennium BP) mummies in Egypt (Keimer 1924; Täckholm 1976). Chemical analysis of Egyptian textiles dated from the twelfth dynasty showed safflower to be one of the dyes used (Darby et al. 1977).This dye plant is also well identified in early Mesopotamian cuneiform records.

C H A PTER 1 0

Plant remains in representative archaeological sites

This chapter summarizes the information on plant remains retrieved from representative Neolithic and Bronze Age sites in west Asia, Europe, and north Africa. It presents a selected list which was compiled in order to answer the question, if one has to sketch the origin and the early spread of domesticated plants in the Old World, what would be the minimum number of archaeological locations that could provide an adequate account on the present state of our knowledge? The information is arranged country by country. Representative sites had to be selected from countries which are still very poorly studied, as well as from areas that had a long tradition of archaeological excavation. In order to attain this goal, numerous well-analyzed locations in intensively researched countries were, not included in this chapter, while poorer sites in less thoroughly surveyed countries do appear on the list. Altogether some 191 sites (or groups of sites) were chosen to represent the archaeological evidence as it stands today. Most sites are radiocarbon-dated, some were dendro-dated; for others we have only estimated dates. Many of the radiocarbon dates are AMS dating, but certainly not all of them. When available, both uncalibrated and calibrated radiocarbon dates were given, using OxCal Program v3.10. In order to compare between sites and strata (since we did not intend to cover the broad issue of dating in full) all dates were rounded to the nearest fifty years. Traditional dating has been given where radiocarbon analyses were not available. If needed, the reader is advised to seek accurate dates and the type of material dated in the cited references. More dates are available in various web-based databases and

online journals, such as ‘C14 radiocarbon CONTEXT’ (http://context-database.uni-koeln.de/index.php), ‘Radiocarbon’ (http://www.radiocarbon.org), or ‘Archaeometry’ (http://www.wiley.com/bw/journal.asp?ref=0003-813x). A general chronological chart (Appendix B) for the different regions is given on pp. 198–199. This book presents archaeological data from sites across vast geographical area and long chronological periods. In the literature, one can find different time periods and geographical boundaries for each region covered. It is beyond the scope of this book to create one sytem that will cover it all. Therefore, hereafter we rely on the terminology used in the archaeobotanical literature cited. However, to make the first stages of agriculture easier to understand, we used one chronological framework for southwest Asia, as Table 9 presents. For information on the geographic location of the sites, consult Appendix A (Maps 21–24, pp. 194–196) and the general references given for each country.

Iran (General reference: Miller 1991; Charles 2007) 1. Ali Kosh, Deh Luran Plain, Khuzistan (Helbaek 1969). (i) Middle PPNB period, Bus Mordeh phase (8450±90 to 8000±50 uncal BP = ca. 9,600–8,750 cal BP). Rich remains: wild-type einkorn wheat (few); einkorn wheat (rare); emmer wheat (prevailing); brittle two-rowed barley (frequent); naked barley (few). Wild: Linum (rare); Prosopis farcta, Pistacia atlantica, and Capparis spinosa (rare). (ii) Middle PPNB period, Ali Kosh phase (ca. 8,290 uncal BP = ca. 9,400–9,250 cal BP). Rich remains: 169

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Table 9 Chronology of cultural entities for southern Levantine Pre-Pottery Neolithic Periods (after Goring-Morris and Belfer-Cohen 2011). Period

Entity/phase

Calibrated 14C years BP

Late Epipaleolithic Pre-Pottery Neolithic A Pre-Pottery Neolithic B

Final Natufian PPNA Early PPNB Middle PPNB Late PPNB Final PPNB/PPNC Yarmukian

12,500–11,750 12,175–11,000 10,950–10,300 10,150–9,725 9,400–8,900 9,050–8,450 8,400–7,700

Early Pottery Neolithic

brittle einkorn wheat (rare); einkorn wheat (rare); emmer wheat (prevailing); two-rowed barley (few); naked barley (rare); lentil (rare). Wild: Linum (rare); Avena (few); Prosopis farcta (frequent); Pistacia atlantica and Capparis spinosa (rare). (iii) Pottery Neolithic, Mohammed Jaffar phase (ca. 7,550 uncal BP = ca. 8,400–8,350 cal BP). Rich remains: emmer wheat (frequent); two-rowed barley (prevailing); sixrowed barley (rare); lentil (rare). Wild: Linum (rare); Avena (frequent); Pistacia atlantica and Capparis spinosa (rare). 2. Tepe Sabz, Deh Luran Plain, Khuzistan (Helbaek 1969). Chalcolithic, Sabz phase and Khazineh phase (ca. 7,450–6,950 uncal BP = ca. 8,350–7,750 cal BP). Scarce remains: wild-type einkorn wheat (few); einkorn wheat (few); emmer wheat (few); free-threshing wheat (frequent); tworowed barley (prevailing); six-rowed barley (frequent); naked barley (few); lentil (frequent); grass pea (few); flax (frequent); Wild: Triticum boeoticum (few); Avena (few); Capparis (few); Prosopis (frequent); Amygdalus (few); Pistacia (few); legumes (frequent); other wild grasses (few). 3. Tepe Hasanlu, Solduz Valley (Tosi 1975). Pottery Neolithic. (i) Hajji Firuz Tepe, periods X–VIII (ca. eighth to the sixth millennia BP). Unspecified quantities: emmer wheat (frequent); two-rowed barley (prevailing). (ii) Pisdeli Tepe, period VIII (ca. sixth millennium BP). Rich remains: emmer wheat (frequent); free-threshing wheat (frequent); two and six-rowed barley (prevailing). 4. Tepe Yahya and adjacent sites, Dowlatabad Plain 200 km south of Kerman (Costantini and Costantini-Biasini 1985). Pottery Neolithic, periods VII and VI (late eighth millennium BP to the seventh millennium BP). Rich remains, unspecified

quantities. The main crops are einkorn wheat, emmer wheat, two and six-rowed barley. Freethreshing wheat and a rounded, small-grained ‘sphaerococcum’ like barley, and broomcorn millet, are also present.

Iraq (General references: Braidwood 1960; Renfrew 1984; Miller 1991; Charles 2007) 1. Jarmo, Kurdistan (Helbaek 1959a, 1960; Braidwood 1960). PPNB (ca. 8,350 uncal BP = ca. 9,450–9,300 cal BP). Scarce remains (not yet fully published, both imprints and carbonized remains): brittle and non-brittle einkorn wheat (rare); brittle and non-brittle emmer wheat (frequent); brittle two-rowed barley (frequent); non-brittle two-rowed barley (few); lentil (rare); pea (rare). Wild: Pistacia; Prosopis; Aegilops; Lathyrus. 2. Tell es-Sawwan, Samarra (Helbaek 1964b). Pottery Neolithic (ca. 7,300–7,000 BP). Numerous remains: einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (few); two-rowed and six-rowed barley (prevailing); naked barley (frequent); flax (rare). Wild: Prosopis farcta (frequent); Capparis spinosa (frequent). 3. Yarym Tepe, northern Iraq (Bakhteyev and Yanushevich 1980). (i) Yarym Tepe I, Pottery Neolithic (eighth millennium BP). Numerous remains: emmer wheat (few); free-threshing wheat (few); spelt wheat? (rare); hulled two-rowed barley (rare); hulled six-rowed barley (prevailing); naked barley (few). (ii) Yarym Tepe II Chalcolithic (sixth millennium BP). Rich remains: emmer wheat (few); free-threshing wheat (rare); spelt wheat? (few); hulled six-rowed barley (prevailing); naked barley

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

(few). No other plants mentioned. (Note that the identification of spelt wheat is based on kernel morphology only. It cannot be regarded as definite.) 4. Choga Mami, Mandali (Helbaek 1972). (i) Samarra phase, Pottery Neolithic (second half of eighth millennium BP). Rich remains: brittle einkorn wheat (rare); einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (frequent); brittle two-rowed barley (frequent); two-rowed and sixrowed barley (rare); naked barley (frequent); lentil (frequent); pea (few); flax (frequent). Wild: Avena sp. (few); Lolium sp. (prevailing); other grasses (frequent); Pistacia atlantica (few). (ii) Post-Samarra phase (ca. 6950 uncal BP = ca. 7800–7750 cal BP). Numerous remains: brittle einkorn and einkorn wheat (rare); emmer wheat (few); free-threshing wheat (rare); six-rowed barley (few); naked barley (rare); lentil (few); flax (rare). Wild: Lolium sp. (prevailing); Pistacia atlantica (few).

Turkey (General references: Nesbitt and Samuel 1996b; Miller 1991) 1. Çayönü, near Ergani, Diyarbakir Province (Van Zeist 1972, 143–66; Van Zeist and De Roller 1991–2, 2003). PPNB (ca. 9,150–8,650 uncal BP = ca. 10,250–9,550 cal BP). Rich charred remains (including chaff): emmer wheat (frequent), the kernels are still narrow resembling those of wild forms, but hundreds of spikelet forks and of glume bases, all have the domestic-type disarticulation scars; pea (frequent), including very few seeds with rough and with smooth seed coat; lentil (frequent); bitter vetch (very frequent); chickpea (rare); flax (few). Wild: Hordeum spontaneum (rare); Secale sp. (rare); Vicia sp. (frequent); Lathyrus cicera/sativus (rare); Linum sp. (few); Ficus carica (rare); Vitis vinifera (few); Pistacia (frequent); Amygdalus sp. (few); Quercus sp. (rare); Celtis sp. (few); several weeds. 2. Can Hasan III, Konya plain (Hillmam 1972, 1978). PPNB (ca. 8,400–7,700 uncal BP = ca. 9,450– 8,450 cal BP). Numerous remains, both charred grains and chaff: einkorn wheat (few), both wild and domestic forms; emmer wheat (frequent); freethreshing Triticum aestivum-like wheat (prevailing), represented by numerous kernels and rachis frag-

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ments; two-rowed hulled barley (frequent); several plump rye kernels; lentil (few); bitter vetch (frequent). Wild: Vicia sativa-like (frequent); Juglans regia (rare); Celtis tournefortii (frequent); Vitis vinifera subsp. sylvestris (rare); Prunus sp. and Crataegus sp. (rare); Lithospermum arvense (very common). 3. Hacilar, Konya Plain (Helbaek 1970). Pottery Neolithic (ca. 7,350–7,000 uncal BP = ca. 8,200–7,800 cal BP). Rich remains: brittle einkorn wheat (frequent); einkorn wheat (frequent); emmer wheat (few); free-threshing wheat (few); two-rowed barley (rare); six-rowed barley (few); naked barley (prevailing); lentil (few); bitter vetch (few). Wild: Pisum sp. (frequent); Aegilops umbellulata (rare); Pistacia atlantica (rare); Celtis australis (frequent); Capparis spinosa (few); Malus sp. (rare); Amygdalus sp. (rare). 4. Çatalhöyük East, Konya Plain (Helbaek 1964a; Fairbairn et al. 2002, 2005, 2007). (i) PPNB, layers Pre-XIIA–D (8,240±55 uncal BP to 8,090±55 uncal BP = ca. 9,350–8,950 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (prevailing); freethreshing wheat (frequent); naked six-rowed barley (rare); rye (few); lentil (prevailing); pea (few); bitter vetch (frequent). Wild einkorn, probable weed (few). Wild fruits and seeds: Celtis (frequent); Pistacia (frequent); Quercus acorns (frequent); Amygdalus sp. (frequent); Malus sp. (rare); Pisum elatius (rare); Ficus (rare); Prunus (rare); Rubus (rare); Capparis (rare); Rhus coriaria (rare). Tubers of Bolboschoenus maritimus (frequent). (ii) Neolithic, Level XII–VI (8,090±55 uncal BP to 7,521±77 uncal BP = ca. 8,950–8,350 cal BP). Rich remains including storage contexts, most assemblage equivalent to Aceramic Neolithic, except: naked six-rowed barley (frequent). Wild fruits and seeds: Celtis (predominant); Juniperus sp. (rare); wild crucifers in storage contexts Capsella bursa-pastoris and Descurainia (frequent); Helianthemum (rare); Taeniatherum caputmedusae (rare); Eremopyrum (rare). Tubers of Bolboschoenus maritimus (few). 5. Aşikli Höyük, Central Anatolia (van Zeist and de Roller 1995). PPNB (8,920±50 to 8,515±40 BP = ca. 10,200–9,500 cal BP). Numerous charred remains (including chaff): einkorn wheat (few); emmer wheat (prevailing); free-threshing wheat (few); hulled two-rowed barley (frequent); naked barley (few); lentil (few); pea (rare); bitter vetch (frequent).

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DOMESTICATION OF PLANTS IN THE OLD WORLD

Wild: Celtis tournefortii (frequent); Pistacia sp. (few); Amygdalus sp. (rare). 6. Erbaba, Beysehir, south-central Anatolia (van Zeist and Buitenhuis 1983). Pottery Neolithic (ca. 5,800–5,400 uncal BC = ca. 8,550–8,150 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat and free-threshing wheat (prevailing); spelt wheat? (rare); hulled two-rowed barley (frequent); naked barley (frequent); lentil (frequent); pea (co-prevailing); bitter vetch (frequent). Wild: Lathyrus cf. cicera; Triticum boeoticum and several herbaceous plants and weeds. 7. Girikihaciyan, near Diyarbakir (van Zeist 1979–80). Halafian (Chalcolithic, ca. 6,950–6,450 uncal BP = ca. 7,800–7,350 cal BP). Numerous remains: einkorn wheat (rare); emmer wheat (prevailing); free-threshing wheat (rare); two-rowed barley (few); lentil (frequent); chickpea (few); bitter vetch (frequent); flax (few). Wild: Pistacia atlantica (rare); Amygdalus sp. and Crataegus sp. (rare); Vicia sp. (few); Lolium sp. (few). 8. Troy and adjacent Kumtepe, the Troad, northwest Turkey (Riehl 1997). (i) Kumtepe, Neolithic Period (ca. 4,805–4,370 cal BC = ca. 6,750–6,300 cal BP). Few charred remains: einkorn wheat, grains and chaff (rare); emmer wheat, chaff (rare); sixrowed hulled barley, grains and chaff (rare); lentil (frequent); bitter vetch (prevailing); flax (rare); fig (frequent). (ii) Kumtepe, Chalcolithic Period (ca. 3,370–2,910 cal BC = ca. 5,300–4,850 cal BP). Few charred remains: einkorn wheat, chaff and grains (frequent); emmer wheat, chaff, and grains (prevailing); six-rowed hulled barley, grains, and chaff (few); lentil (rare); faba bean (rare); bitter vetch (few); Lathyrus cicera/sativus (rare); fig (few); grapevine (few). (iii) Troy, Early Bronze Age (ca. 2,900– 2,200 BC = ca. 4,850–4,200 cal BP). Few charred remains: einkorn wheat, chaff, and grains (frequent); emmer wheat, grains, and chaff (prevailing); six-rowed hulled barley, grains, and chaff (few); lentil (rare); bitter vetch (few); Lathyrus cicera/sativus (rare); olive (rare); fig (few); grapevine (few). (iv) Troy, Middle Bronze Age (ca. 2,200–1,700 BC = ca. 4,150–3,650 cal BP). Very rich charred remains: einkorn wheat, chaff, and grains (few); emmer wheat, chaff, and grains (frequent); freethreshing hexaploid wheat, chaff, and grains (few); six-rowed hulled barley, grains, and chaff (fre-

quent); lentil (rare); pea (frequent); bitter vetch (few); Lathyrus cicera/sativus (rare); faba bean (rare); flax (prevailing); fig (few); grapevine (few); gold of pleasure (few). (v) Troy, Late Bronze Age (ca. 1,700– 1,200 BC = ca. 3,650–3,200 cal BP). Rich charred remains: einkorn wheat, grains, and chaff (few); emmer wheat, chaff, and grains (few); free-threshing hexaploid wheat, chaff, and grains (few); sixrowed hulled barley, grains, and chaff (frequent); broomcorn millet (rare); lentil (rare); chickpea (prevailing); bitter vetch (frequent); Lathyrus cicera/sativus (rare); faba chickpea (rare); flax (prevailing); fig (rare); grapevine (rare); gold of pleasure (rare).

Syria (General reference: Miller 1991; Willcox 2007; Willcox et al. 2008, 2009) 1. Tell Abu Hureyra, Northern Syria, Euphrates valley (Hillman 1975, 1989, 2000a, 2000b; De Moulins 1997, 2000; Hillman et al. 2001). (i) EpiPalaeolithic (ca. 12,700–11,100 cal BP). Rich remains of charred seeds belonging to a wide range of species, all conforming to wild forms in their morphology, and no chaff remains: narrow grains of wild two-grained einkorn wheat (rare) and wild rye (prevailing); small seeded lentil (few); bitter vetch (rare); Pistacia sp. (frequent); Celtis sp. (rare); Capparis spinosa (rare); large numbers of Polygonum and Bolboschoenus maritimus nutlets; seeds of numerous grasses (Stipa prevailing), and herbs (Brassicaceae prevailing). (ii) PPNB culture, four excavation trenches of various strata (from 8640±100 uncal BP, from trench B phase 2 = ca. 9,800–9,500 cal BP, to 802±100 uncal. BP from trench G phase 6 = ca. 9,050–8,700 cal BP). Numerous, various charred remains from the four trenches in moderate quantities in each sample: wild and domesticated einkorn, grain and chaff; hexaploid or tetraploid wheat; wild and domesticated barley; wild and domesticated rye; lentil; chickpea; large vetch; Prosopis farcta; small legumes (frequent); Capparis spinosa; grape pips (few). Many other grass and herb seeds and uncharred Boraginaceae. 2. Tell Mureybit, northern Syria, Euphrates valley (van Zeist and Casparie 1968; van Zeist and Bakker-Heeres 1986). Epi-Palaeolithic (ca. 10,200–

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

9,900 uncal BP = ca. 11,800–11,300 cal BP). Rich remains of charred seeds—all conforming in their morphology to wild forms: narrow grains of wild two-grained einkorn wheat and of wild rye (prevailing); glumes of rye (few); narrow, wild-type grains of barley (few); small-seeded lentil (few); bitter vetch (rare); wild flax (rare); Pistacia atlantica (rare); numerous grass and herb seeds. 3. Jerf el Ahmar, northern Syria, Euphrates valley (Willcox 2000; Willcox et al. 2008, 2009). (i) PPNA early phase (ca. 9,800–9,500 uncal BP = ca. 11,500– 11,000 cal BP). Rich charred remains, chaff indicating that all cereals conform to wild forms in their morphology: two-grained wild einkorn, T. boeoticum/T. urartu (rare); wild rye, Secale sp. (prevailing); wild barley (prevailing); Lens sp. (frequent); Vicia ervilia (rare); wild small grasses (H. murinum type, Stipa sp., Taeniatherum) (frequent). Pistacia sp. (frequent); Amygdalus sp. (frequent); Polygonum/Rumex (rare).1 (ii) PPNA late phase (ca. 9500–9300 uncal BP = ca. 11,000–10,600 cal BP). Similar to the early phase, only wild barley increases and wild rye declines. 4. Djade el Mughara, northern Syria, Euphrates valley (Willcox et al. 2008). PPNA (ca. 9,300–9,000 uncal BP = ca. 10,700–10,400 cal BP). Similar to Jerf el Ahmar, except that wild-type founder species, emmer, single-grained einkorn, chickpea, and faba bean appear for the first time in northern Syria. 5. Tell Aswad, south-east of Damascus (van Zeist and Bakker-Heeres 1985). (i) Early PPNB (ca. 9,300– 9,000 uncal BP = ca. 10,500–10,200 cal BP). The dates of these levels (considered initially to be PPNA) have been recently revised based on AMS dating of emmer grains (Willcox 2005, Table 1) retrieved from the early excavations studied by van Zeist, as well as more recent excavations. Numerous remains of charred seeds: plump domestic-type grains of emmer wheat (frequent); either wild or domestic two-rowed barley (few); small-seeded lentil (rare); pea (few); Pistacia sp. (prevailing); Ficus carica (frequent); Capparis spinosa (rare); Amygdalus sp. shells (rare); Trigonella sp. (frequent); seeds of numerous grass and herb species. (ii) Middle PPNB (ca. 10,200–9,550 cal BP). Numerous remains of charred seeds: plump, domestic-type grains of emmer wheat (frequent); free-threshing 1

Willcox argues for pre-domestic agriculture.

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wheat (few); either wild or domestic two-rowed barley (rare); naked barley (rare); small-seeded lentil (rare); pea (few); early finds of flax (frequent). Wild: Pistacia sp. (frequent); Ficus carica (frequent); Capparis spinosa (rare); Vitis vinifera (rare); Trigonella sp. (frequent); Cyperus sp., Carex sp., Lolium sp., and Phalaris sp. (frequent); numerous grass and herb seeds.

Israel and Jordan (General references: Miller 1991; Neef 1998) 1. Ohalo II, Sea of Galilee (Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004, 2008). Upper Palaeolithic (ca. 23,000 cal BP). Rich charred remains, ca. 140 wild taxa: wild emmer wheat, T. dicoccoides (frequent); wild barley, H. spontaneum (prevailing); wild lentil, L. orientalis (rare); wild pea, Pisum sativum subsp. humile (rare). Nine edible wild fruits: wild almond, Amygdalus communis (rare); Crataegus aaronia/azarolus (few); Nitraria schoberi (frequent); Olea europaea (rare); Pistacia atlantica (rare); Pyrus syriaca (rare); Quercus sp. (few); Vitis vinifera (rare); Zizyphus spina-christi (rare). Grains of several wild grasses such as Aegilops, Avena, and Bromus; seeds of several wild herbs. 2. Netiv Hagdud and Gilgal, Lower Jordan Valley (Kislev 1997; Hartmann 2006; Weiss et al. 2006). PrePottery Neolithic A (ca. 10,000–9,400 uncal BP = ca. 11,700–10,550 cal BP). Rich remains of charred wild plant remains: dicoccoides-type emmer wheat (frequent); spontaneum-type barley (prevailing, storage); other wild Horduem species (few); Avena sterilis (frequent, storage); A. wiestii (rare); Aegilops sp. (rare); Alopecurus utriculatus (few); Lens sp. (frequent); other legume species (rare); Ficus carica (very common); Vitis vinifera (rare); Pistacia atlantica (few); Quercus ithaburensis (rare); Amygdalus communis (rare). 3. Nahal Zehora II , Mount Carmel , Israel (Kislev and Hartmann forthcoming). Pottery Neolithic (ca. 7,300–7,100 uncal BP = ca. 8,150– 7,850 cal BP). Rich charred remains: emmer wheat (prevailing); parvicoccum-type wheat (frequent); barley (few); lentil (few); faba bean (rare); flax (rare). Fruit trees: fig (rare); olive (rare); Pistacia atlantica/palaestina (rare). Wild/weedy grasses, including Lolium temulentum (rare).

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4. Jericho, Lower Jordan Valley (Hopf 1983). (i) Middle Pre-Pottery Neolithic B (ca. 9,900–9,550 cal BP). Rich remains: einkorn wheat and emmer wheat (prevailing); two-rowed barley (frequent); lentil (frequent); pea (frequent); Vicia faba-type (few); chickpea (rare); flax (an imprint of a single capsule). (ii) Ceramic Neolithic (first half of the seventh millennium BP). Rich remains: einkorn wheat and emmer wheat (frequent); two-rowed barley (prevailing). Wild: some grass and herb seeds. (iii) Chalcolithic (early sixth millennium BP). Numerous remains: wheat (rare); two-rowed barley (prevailing). Wild: Ficus carica (rare). (iv) Early Bronze Age (ca. 3,500–3,300 BPBP). Rich remains: einkorn wheat (frequent); emmer wheat (prevailing); free-threshing wheat (frequent); two-rowed and six-rowed barley (frequent); lentil (frequent); chickpea (few); flax, a capsule and seed (rare); grapevine (frequent, both pips and berries); date palm (rare); fig (rare). Wild: Allium cf. ampeloprasum (few); Pistacia atlantica (rare); numerous grass and herb seeds. (v) Middle Bronze Age. Rich remains: emmer wheat (frequent); free-threshing wheat (frequent); six-rowed barley (prevailing); lentil (rare); pomegranate (rare). Wild: Allium cf. ampeloprasum (rare); some grass and herb seeds. 5. Atlit Yam, Carmel Coast (Kislev et al. 2004; and pers. comm.). Pre-Pottery Neolithic C (ca. 8,000– 7,500 cal BPBP). Rich charred and waterlogged remains in a well: emmer wheat (prevailing); naked (T. parvicoccum) wheat (frequent); barley (few); lentil (few); chickpea (rare); flax. Fruits and nuts: Ficus carica (frequent); Vitis sylvestris; Amygdalus communis/korschinskii; Pistacia atlantica; P. lentiscus; P. Palaestina; Phoenix theophrasti. Large group of wild and weedy plants: Papaver setigerum; Cuminum cyminum; Pinus halepensis; Lolium temulentum; Phalaris paradoxa; Mercurialis annua; Malva parviflora; Rumex pulcher; Daucus carota; Coriandrum sativum; Murinum sylibum; Capparis spinosa; Rubus sanguineus. 6. Tuleilat Ghassul, Lower Jordan Valley and Shiqmim, Negev (Zohary and Spiegel-Roy 1975; Kislev 1987). Late Chalcolithic (ca. 6,800–5,800 cal BP). Rich remains: emmer wheat (few); two and sixrowed barley (frequent); lentil (few); olive (frequent, in Ghassul only); date palm (few, in Ghassul only). 7. Tell es-Sa’idiyeh, Jordan (Cartwright 2003; and pers. comm.). Early Bronze Age (ca. 4,900 BP).

Very rich charred remains, in a palace’s kitchen: wheat; barley; lentil; chickpea; faba bean-type; grapes; figs; pomegranate; olive; Wild: Cratagus sp.; Ziziphus spina-christi; Pistasia sp.; Quercus sp.; Capparis spinosa, flower buds. 8. Bab edh-Dhra, south-east of Dead Sea (McCreery 1979). Early Bronze Age. Rich remains: emmer wheat (probably also einkorn and freethreshing wheat) (few); two-rowed barley (few); six-rowed barley (prevailing); lentil (few); chickpea (frequent); linseed (few); olive (frequent); grapevine (frequent, whole berries and pips); fig (frequent); almond (rare). Wild: Pistacia atlantica (few); Prunus insititia-like (rare); Lolium temulentum and herb seeds. Textiles made of flax fibres.

Egypt (General references: Täckholm 1976; Germer 1985; Wetterstrom 1993, 1998; Murray 2000a, 2000b, 2000c; de Vartavan et al. 2010). 1. Nabta Playa, Western Desert, south Egypt (Wasylikowa 1997; Wasylikowa et al. 1997; Wasylikowa and Dahlberg 1999). Site E-75–6. Preagriculture site (8,095±12 to 7,950±90 uncal BP = ca. 9,250–8,650 cal BP). Site E-75–6. Rich charred remains of wild plants: Sorghum bicolor subsp. arundinaceum (very frequent); Echinochloa colona (very frequent); Panicum turgidum (frequent); several other grass species (few); Arnebia sp. (frequent); Capparis cf. decidua (few); Schouwia purpurea (very frequent); Boerhavia sp. (frequent); several legumes (very frequent); Zizyphus cf. spina-christi (frequent); tubers of Cyperus cf. rotundus (frequent). 2. Farafra Oasis, Hidden Valley, Western Desert (Barakat and Fahmy 1999; Fahmy 2001). Neolithic period (6,028±150 to 5,163±374 uncal BC = ca. 7,050– 5,450 cal BP). Few charred remains of wild plants, grasses predominant: Sorghum sp. (prevailing); Setaria verticillata (frequent); Panicum repens (frequent); Echninochloa colona (frequent); Brachiaria sp. (few); Cenchrus (rare); Digitaria sp. (rare); Citrullus colocynthis (rare). 3. Maadi, near Cairo (van Zeist and de Roller 1993; van Zeist et al. 2003). Late Neolithic, Predynastic (4,680±70 to 4,900±70 uncal BP = ca. 5,650–5,450 cal BP). Rich charred and desiccated remains: emmer wheat, grain and chaff (prevailing); two-rowed and

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

six-rowed barley (frequent); lentil (rare); linseed (rare); Ficus sp. (rare); Cucumis cf. melo (rare). 4. Fayum, Kom K site (Caton-Thompson and Gardner 1934; Wetterstrom 1993; Wendrich and Cappers 2005). Early Neolithic (6,095±250 to 6,391±180 uncal BP = ca. 7,500–6,650 cal BP). Large quantities of parched and of charred grains in underground silos: mainly emmer wheat, tworowed, and six-rowed barley; also flax textile. 5. Merimde, Beni Salâme, western Nile Delta. (i) Neolithic (6,130±110 BP to 5,260±90 uncal BP = ca. 7,150–5,950 cal BP) (Wetterstrom 1993). Rich remains: emmer wheat (prevailing); free-threshing wheat? (rare); hulled six-rowed barley (frequent); lentil (few); pea (few); flax (rare). Wild: Lolium and several other weeds, sedges, and legumes (frequent). (ii) Middle Neolithic (Helbaek 1955). Rich remains: emmer wheat (prevailing); free-threshing wheat (rare and later absent); hulled six-rowed barley (frequent). 6. Naqada (Nagada) area, Upper Egypt (Wetterstrom 1993). (i) Site KH3, Nagada I (ca. 5,900–5,650 BP). Numerous remains: emmer wheat (prevailing); hulled six-rowed barley (frequent); pea (rare); flax (rare). Wild: Citrullus colocynthis (rare); Zizyphus spina-christi (rare). (ii) South Town, Nagada II (ca. 5,650–5,300 BP). Numerous remains: emmer wheat (prevailing); hulled six-rowed barley (frequent); pea (rare); bitter vetch (rare); flax (few). Wild: Citrullus colocynthis (few); Zizyphus spinachristi (rare); some weeds. 7. Hierakonpolis, Upper Egypt (Fahmy 1995, 2003, 2005; Fahmy et al. 2008). (i) Site HK11C, trash mound, Naqada IIAB (ca. 5,700–5,600 BP). Rich remains: emmer wheat (prevailing); free-threshing wheat (rare); hulled six-rowed barley (frequent); hulled two-rowed barley (rare); naked barley (rare); Cucumis melo (frequent); flax (frequent). Wild: Balanites aegyptiaca (rare); Ceruana pratensis (frequent); Fimbristylis bisumbellata (rare); Setaria verticillata (rare). Weeds: Lolium temulentum (few); Phalaris minor (frequent); as well as other field weeds. (ii) Site HK43, Workers’ Cemetery, Naqada IIBC (ca. 5,450–5,300 BP). Numerous remains: emmer wheat (prevailing); free-threshing wheat (rare); hulled six rowed barley (frequent); hulled two-rowed barley (rare); naked barley (rare); Cucumis melo (rare). Wild: large quantities of culms

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of Juncus, which have been used for making of matting; Anethum graveolens (frequent); Balanites aegyptiaca (rare); Ceruana pratensis (rare); Cyperus esculentus (rare); Acacia (frequent); Ziziphus spinachristi (frequent); also flax textile. 8. Kom el-Hisn, west part of the Nile delta (Moens and Wetterstrom 1988). Old Kingdom deposits (ca. 4,700–4,250 BP). Numerous remains: emmer wheat (prevailing); barley (frequent); pea (rare); flax (rare); clover seeds, probably Trifolium alexandrinum (co-prevailing). Wild: Lolium temulentum and Phalaris paradoxa (frequent); numerous herbs, reeds, and sedges. 9. Saqqara, Memphis. (i) Djoser (Zoser) pyramid, thirdrd dynasty, ca. 4,630–4,611 BP (Lauer et al. 1950). Rich remains (desiccated material): emmer wheat and hulled six-rowed barley (the bulk of the find); lentil (rare); sycamore fig (numerous dry sycons). Apparently imported: fragments of grapevine raisins (few); Mimusops schimperi (rare); Juniperus oxycedrus (few); Wild: Lolium temulentum (the commonest weed); Phalaris paradoxa (rare); Vicia sativa (few); V. lutea (few); V. narbonensis (rare); Lathyrus aphaca (few); L. marmoratus (rare); L. hirsutus (few); Scorpiurus muricata (few); Trigonella hamosa (rare); Medicago hispida (rare); Rumex dentatus (few); Anthemis pseudocotula (rare); Beta vulgaris (rare); Zizyphus spina-christi (numerous); Acacia nilotica (rare); Balanites aegyptiaca (rare). (ii) Queen Icheti’s tomb, sixth dynasty, ca. 4,550 BP (Helbaek 1953). Rich remains (mummified material): emmer wheat prevailing. 10. Tutankhamun tomb, Valley of the Kings, near Thebes (Germer 1989a; Hepper 1990; de Vartavan et al. 2010) New Kingdom, nineteenth dynasty (ca. 1,325 BC). An extremely rich find of desiccated, excellently preserved plants and plant products: emmer wheat (main element in the model granary); six-rowed barley (large quantities, in baskets, including germinated seeds in ‘Osiris bed’); lentil (few); chickpea (few); fenugreek (in several baskets, mixed with coriander); flax (linen, bow strings, few seeds); olives (leaves and twigs in garlands, jars of olive oil); grape wine (shrivelled berries and their pips, wine amphoras with records on production sites and dates); sycamore fig (fruits and timber); date palm (fruits in basket, ropes); almond (basket with fruits);

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DOMESTICATION OF PLANTS IN THE OLD WORLD

watermelon (numerous seeds in basket); garlic (numerous bulbs, some with leaves); black cumin (numerous seeds); coriander seeds (main item in several baskets); safflower (numerous achens, textiles dyed with safflower pigment). Remains of several ornamental plants and plants collected from the wild (Grewia sp.; Ziziphus spina-christi; Juniperus sp.). Timber (in furniture and implements) of numerous local and foreign trees, including imported cedar (Cedrus libani) and ebony (Dalbergia melanoxylon).

Libya (General reference: van der Veen 1995, 2006) 1. Zinchecra, Hilltop settlement, Fazzan, southern Libya (van der Veen 1992a, 1992b). Early Garamantian (ca. 2,700–2,400 cal BP). Rich charred remains: emmer wheat (frequent); bread wheat (rare); six-rowed barley (frequent); date palm (frequent); grapevine (rare); fig. Herbs: celery (Apium graveolens); purslane (Portulaca oleracea); dill (Anethum graveolens); fennel (Foeniculum vulgare). Wild: Citrullus colocynthis; Rhus tripartita. 2. Jerma (Garama), Wadi settlement, Fazzan, southern Libya (Pelling 2008). (i) ca. twenty-fourth to the twenty-first century BP, Scarce remains: emmer wheat (rare); barley (rare); fig (rare); grapevine (rare); date palm (rare). Wild: Pennesitum glaucum. (ii) ca. 2,200–1,600 cal BP, rich charred remains: durum wheat; bread wheat; barley; sorghum (Sorghum bicolor); olive; grapevine; fig; date palm; almond; pomegranate; cotton (Gossypium sp.). Wild: Pennisetum glaucum.

emmer wheat; free-threshing wheat; hulled barley; naked barley; several wild grasses and trees. 2. Arukhlo 1 and Arukhlo 2, Bolnisskiy region, Georgia (Januševič 1984; Lisitsina 1984; SchultzeMotel 1988). Neolithic, Aruchlo 1 Period (7,135±70 uncal BP to 6,365±140 uncal BP = ca. 8,000–7,150 cal BP). Unspecified quantities (except for the wheats), mostly imprints: einkorn wheat (few); emmer wheat (numerous); free-threshing wheat (prevailing); spelt wheat (rare); two-rowed and six-rowed hulled barley; naked barley; broomcorn millet; lentil; pea; bitter vetch. 3. Imiris-Gora, Marnsulskij region, Georgia (Lisitsina and Prishchepenko 1977; Schultze-Motel 1988a). Eneolithic (ca. 6,300±12 uncal BP = ca. 7,400– 7,000 cal BP). Imprints, unspecified quantities: emmer wheat; free-threshing wheat; spelt wheat(?); six-rowed hulled and naked barley; broomcorn millet(?); Avena sp. 4. Aratashen and Aknashen, Ararat Valley, Armenia (Hovsepyan and Willcox 2008). Neolithic culture (ca. 7,035±69 uncal BP to ca. 6,350±70 uncal BP = ca. 7,950–7,150 cal BP). Numerous charred remains and imprints: einkorn wheat (rare); emmer wheat (few); free-threshing wheat (frequent); hulled barley (rare); naked six-row barley (frequent); smallseeded lentil, seeds and pods (frequent); bitter vetch (few). Wild: Alyssum desertorum (prevailing, capsule imprints); Camelina microcarpa (frequent, capsule imprints); Vitis sylvestris; Celtis sp.; Elaeagnus sp.; Buglossoides arvensis (frequent); Rumex cf. crispus; Bolboschoenus maritimus.

Central Asia Morocco Kaf That el-Ghar, NW Morocco (Ballouche and Marinval 2003). Early Neolithic, Cardial culture (ca. 6,500–5,500 cal BP). Few charred remains: emmer wheat (prevailing); bread/durum wheat (few); naked barley (very rare); broad bean (rare).

Caucasia and Transcaucasia (General reference: Wasylikowa et al. 1991) 1. Chokh, Dagestan (Lisitsina 1984). Neolithic (end of eighth millennium BP to the seventh millennium BP). Unspecified quantities: einkorn wheat;

(General reference: Harris and Gosden 1996) 1. Jeitun (Djeitun), south Turkmenistan (Charles and Hillman 1992; Harris et al. 1993; Harris et al. 1996). Neolithic Jeitun culture (7,270±100 to 7,100±90 uncal BP = ca. 8,200–7,850 cal BP). Numerous charred remains: einkorn wheat (prevailing); emmer wheat (few); six-row barley (few). Rare trace of freethreshing wheat (?). 2. Anau, south Turkmenistan (Miller 1999; Miller, 2003). Chalcolithic (Anau North, ca. 6,500–5,000 cal BP) and Bronze Age (Anau South, ca. 5,000–3,700 cal BP). Few charred remains: bread wheat (frequent); six-row barley (prevailing); pea, in Bronze

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

Age only (rare); grape, in Bronze Age only (rare). Wild: Celtis sp.; Capparis sp.; as well as genera from several families notably the Chenopodiaceae, Fabaceae, and Poaceae. 3. Shortugai, Oxus River Valley, Afghanistan (Willcox 1991). Mature Harappan culture (2,245±100 uncal B.C. = 2,350–2,100 cal BP) Numerous remains: free-threshing wheat (frequent); barley (prevailing); broomcorn millet (frequent); lentil (few); pea (few); flax (rare); grapevine (frequent, probably cultivated). Wild: Pistacia vera; Eleagnus angustifolia; Amygdalus sp.; Prosopis sp.; several grasses, including Aegilops tauschii and herbs.

Cyprus (General references: Hansen 1991a, 1994; Kroll 1991; Miller 1991) 1. Kissonerga-Mylouthkia and Shillourokam bos, Cyprus (Willcox 2000; Murray, 2003). CyproEarly PPNB (9,310±80 to 8,670±80 uncal BP = ca. 10,650–9,550 cal BP). Numerous charred and impression remains: einkorn wheat (?), grains, and chaff (rare); emmer wheat (?), grains, and chaff (rare); hulled barley (?) (frequent). Most other taxa were not identified to species level and probably wildgrowing: legumes, fruits, and numerous wild/ weeds taxa. The domesticated status of the cereals requires further support. 2. Cap Andreas-Kastros (van Zeist 1981). Aceramic Neolithic (6,140±200 to 7,775±125 uncal BP = 8,700–6,800 cal BP). Numerous remains: einkorn wheat (frequent); emmer wheat (prevailing); six-rowed barley (frequent); lentil (frequent); pea (?) (rare); flax (rare). Wild: Vicia faba/narbonensis (rare); Olea europaea (rare); Pistacia atlantica/terebinthus (rare); Ficus carica (rare); Lolium sp. (frequent); Vicia sp. (few); Malva sp. (few). 3. Khirokitia (Waines and Stanley Price 1975– 1977; Hansen 1991a, 1994). Aceramic Neolithic (ca. 7500 uncal BP = ca. 8,400–8,200 cal BP). Rich remains (including chaff): einkorn wheat and emmer wheat (prevailing); six-rowed barley (few); lentil (frequent); pea (few). Wild: Lathyrus cf. sativus (rare); Vicia faba/ narbonensis (rare); Ficus carica (frequent); Olea europaea (rare); Prunus insititia (few); Pistacia sp. (rare). 4. Dhali Agridhi, Idalion (Stewart 1974). Aceramic Neolithic (7,990±80 to 7,290±465 uncal BP = ca.

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9,000–7,600 cal BP). Numerous remains: emmer wheat (few); free-threshing wheat (rare); two- and six-rowed hulled barley (frequent); naked barley (few); lentil (frequent); pea (rare). Wild: Olea europaea (frequent); Vitis vinifera (few); Ficus carica (frequent); Amygdalus sp. (rare); Pistacia sp. (rare); Anchusa sp. (frequent). 5. Hala Sultan Tekke, near Larnaca (Hjelmqvist 1979a). Late Bronze Age (ca. 3,200 uncal BP = ca. 3,450–3,350 cal BP). Rich remains: emmer wheat (few imprints); free-threshing wheat (rare); hulled and naked barley (prevailing-imprints); lentil (rare); olive (few); grapevine pips (frequent); fig pips (frequent); pomegranate (rare); citron (rare). Wild: Pistacia terebinthus (rare); Capparis spinosa (rare); Zizyphus spina-christi (rare); Lupinus albus (rare); Chrozophora verbascifolia (few); Onopordon cf. illyricum (few).

Greece (General references: Renfrew 1979; Kroll 1991; Valamoti and Kotsakis 2007). 1. Franchthi Cave, Argolis (Hansen 1991a, 1992). (i) Upper Palaeolithic (ca. 13,000–9,000 uncal BP = ca. 15,500–10,150 cal BP). Scarce remains: Wild: brittle two-rowed barley (few); Lens sp. (few) Avena sp. (rare); Vicia sp. (few); Pistacia sp. (few); Amygdalus sp. (few); Lithospermum, Alkanna, Anchusa (frequent). (ii) Mesolithic (ca. 9,300–8,000 uncal BP = ca. 10,600–8,750 cal BP). Scarce remains: Wild: brittle two-rowed barley (few); Lens sp. (few); Pisum sp. (few); Avena sp. (few); Pistacia sp., Amygdalus sp. Pyrus sp. (few); Lithospermum, Alkanna and Anchusa (frequent). (iii) Early Neolithic (7,794±140 to 6,940±90 uncal BP = ca. 8,800–7,700 cal BP). Scarce remains: emmer wheat (few); two-rowed barley (few). Wild: Pistacia sp. (few); Amygdalus sp. (rare); Lithospermum, Alkanna, Anchusa, Lens sp., Vicia sp. (few). (iv) Middle Neolithic (ca. 7,000–6,300 uncal BP = 7,950–7,200 cal BP). Scarce remains: einkorn wheat (rare); emmer wheat (frequent); two-rowed barley (frequent); lentil (few); bitter vetch (rare). Wild: Pistacia sp. and Amygdalus sp. and Pyrus sp. (rare); Lithospermum, Alkanna, Anchusa (rare). (v) Late Neolithic (ca. 6,300–4,800 uncal BP = 7,300– 5,450 cal BP). Scarce remains: einkorn wheat (frequent); two-rowed barley (frequent); lentil (few);

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bitter vetch (rare). Wild: Pistacia sp. Pyrus sp., Amygdalus sp., Vitis sylvestris (rare); Lithospermum, Alkanna, and Anchusa (rare). 2. Sesklo (including Argissa and Otzaki), Thessaly (Hopf 1962; Kroll 1981a). (i) Aceramic and Early Neolithic (7,755±97 to 7,483±72 uncal BP = ca. 8,650–8,200 cal BP). Numerous remains: einkorn wheat (frequent); emmer wheat (prevailing); sixrowed barley (frequent); lentil (frequent); bitter vetch (few). Wild: Ficus carica (frequent); Vitis vinifera (rare); Sambucus ebulus (rare); Pistacia atlantica (frequent); Lithospermum arvense (frequent); Avena sp. (frequent). (ii) Proto- and pre-Sesklo phases (eighth millennium BP). Rich remains: einkorn and emmer wheat (prevailing); free-threshing wheat (rare); two-rowed barley (few); six-rowed hulled and naked barley (frequent); lentil (frequent); pea (few); bitter vetch (few); chickpea (rare); flax (few). Wild: Ficus carica (frequent); Sambucus ebulus (rare); Pistacia atlantica (rare); Lithospermum arvense (frequent); Avena sp. (few). (iii) Sesklo phase (seventh millennium BP). Numerous remains: einkorn wheat (frequent); emmer wheat (prevailing); six-rowed barley (few); broomcorn millet (rare); lentil (frequent); pea (rare); bitter vetch (rare); flax (rare). Wild: Ficus carica (frequent); Vitis vinifera (rare); Pistacia atlantica (rare); Lithospermum arvense (frequent); Avena sp. (few); (iv) Dimini phase, Late Neolithic (early to middle of sixth millennium BP). Numerous remains; einkorn wheat (frequent); emmer wheat (prevailing); free-threshing wheat (rare); six-rowed barley (frequent); lentil (few); fabalike bean (rare); bitter vetch (rare); flax (rare). Wild: Ficus carica (frequent); Vitis vinifera (frequent); Amygdalus sp. (rare); Lithospermum arvernse (rare); Avena sp. (few). (v) Rachmani phase, Late Neolithic (late sixth to early third millennium BP). Rich remains: einkorn wheat, emmer wheat, six-rowed barley, and lentil (equally frequent); free-threshing wheat (rare); naked barley (rare); pea (rare); grass pea (few); flax (rare). Wild: Ficus carica (frequent); Vitis vinifera (rare); Quercus acorns (rare); Amygdalus sp. (rare); Camelina sativa (a single seed); Lithospermum arvense (few); Avena sp. (few). 3. Nea Nikomedeia, Macedonia (van Zeist and Bottema 1971). Early Neolithic (ca. 7,470 uncal BP = ca. 8,400–8,100 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (prevailing); naked barley

(frequent); lentil (frequent); pea (frequent); bitter vetch (frequent). Wild: Quercus acorns (few); Cornus mas (rare); Prunus cf. spinosa (rare). 4. Toumba Balomenou, Chaeronia, Boiotia (Sarpaki 1995). Numerous remains, mostly fragmented seeds, and chaff. (i) Early Neolithic (ca. 6,800–6,400 BP = ca. 7,700–7,250 cal BP): einkorn wheat, mainly chaff (prevailing); barley (few); lentil (frequent); pea (rare); bitter vetch (rare). Wild: Lolium temulentum; Lathyrus cicera/sativus; Rubus sp.; Pistacia terebinthus; Vitis vinifera; Ficus carica; numerous herbs. (ii) Early-Middle Neolithic (ca. 6,400– 5,600 BP = ca. 7,400–6,300 cal BP): einkorn wheat, mainly chaff (prevailing); barley (few); lentil (frequent); pea (rare); bitter vetch (rare). Wild: Lolium temulentum; Avena sp.; Lathyrus cicera/sativus; Rubus sp.; Pistacia terebinthus; Vitis vinifera; Ficus carica; numerous herbs. 5. Dimini, Thessaly (Kroll 1979). Late Neolithic, Classic phase (5,630±150 uncal BP = ca. 6,650–6,300 cal BP). Rich remains: einkorn wheat (rare); emmer wheat (frequent); six-rowed barley (rare); naked barley (frequent); lentil (frequent); pea (frequent); faba bean (few); bitter vetch (few); chickpea (few); grass pea (frequent). Wild: Vitis vinifera (rare); Amygdalus sp. (few). 6. Lerna, Argolis (Hopf 1961a). (i) Final Neolithic: Ficus carica, Arbutus. (ii) Early Bronze Age (fifth millennium BP). Numerous remains: einkorn and emmer wheats (few); six-rowed barley and naked barley (few); lentil (frequent); pea (few); faba bean (frequent); bitter vetch (frequent); grass pea (?) (few); flax (frequent); fig (frequent, both fruits and pips); grapevine (few). 7. Kastanas, Macedonia (Kroll 1983, 1984). (i) Early Bronze Age (second half of fifth millennium BP). Rich remains: einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (rare); spelt wheat (rare); six-rowed barley (prevailing); lentil (frequent); pea (rare); bitter vetch (frequent); faba bean (rare); grass pea (rare); flax (few); poppy (rare); grapevine (few); fig (frequent). Wild: Quercus acorns (few); Pyrus sp. (few); Rubus fruticosus (rare); Sambucus ebulus (few); Cornus mas (few); Lolium temulentum (few); other grass and herb seeds. (ii) Late Bronze Age (second half of fourth millennium BP). Rich remains: einkorn and emmer wheats (frequent); spelt wheat (rare); free-threshing wheat

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

(rare); six-rowed barley (few); lentil (frequent); bitter vetch (prevailing); broomcorn millet (frequent); foxtail millet (few); poppy (rare); grapevine (frequent); fig (few). Wild: Quercus acorns (rare); Sambucus ebulus (rare); Secale cereale and Avena sp. (contaminating wheat and barley); Camelina sativa (rare); Fragaria sp. (rare); numerous grass and herb seeds.

Crete Knossos, Crete (Sarpaki 2009). (i) Aceramic Neolithic (7,740±130 uncal BP = ca. 8,650–8,400 cal BP). Rich charred remains: einkorn wheat (rare); emmer wheat (rare); bread wheat (prevailing); hulled two-rowed and six-rowed barley (rare); naked six-rowed barley (rare); lentil (few); fig (rare); almond (rare). Wild: Avena sp.; Lolium sp. (ii) Early Neolithic I (ca. 7,200–7,000 uncal BP = ca. 8,050–7,800 cal BP). Few remains, mainly charred: einkorn wheat (rare); ‘eastivo-compactum’ wheat (prevailing); barley (frequent); lentil ? (rare); flax (rare); fig, nutlets and fruits (frequent); almond (frequent). Wild condiments/aromatic: Raphanus raphanistrum (wild radish), pods (frequent); Satureja thymbra (few). (iii) Early Neolithic II (ca. 7,000–6,800 uncal BP = ca. 7,800–7,600 cal BP). Few charred remains: ‘eastivo-compactum’ wheat (frequent); barley (few); lentil (rare); flax? (rare); fig, nutlets and fruits (prevailing); almond (frequent); grapevine (rare). Wild condiments/aromatic: Raphanus raphanistrum (wild radish), pods and seeds (frequent); Satureja thymbra (rare); Thymelaea hirsuta (rare). (iv) Middle and Late Neolithic. Rare remains, mainly charred: wheat/barley (rare); lentil (rare); pea? (rare); fig (frequent); almond (prevailing).

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8,200–6,650 cal BP). Numerous finds: einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (few); six-rowed barley (few); lentil (rare); pea (rare). Wild: Malus sp. Corylus avellana, Cornus mas, and Vitis vinifera (few). (ii) Late Neolithic (ca. 6,100 BP). Scarce remains: einkorn and emmer wheat (frequent); six-rowed barley (rare); lentil (rare); pea (rare). 3. Obre, Bosnia-Hercegovina (Renfrew 1974). (i) Early Neolithic, Starčevo group (7,240±60 to 6,150±60 uncal BP = ca. 8,150–7,000 cal BP). Scarce remains: einkorn wheat (rare); emmer wheat (prevailing); free-threshing wheat (rare); lentil (few); pea (frequent, in a single context). Wild: Cornus mas (rare). (ii) Late Neolithic, Butmir culture (4,000– 3,860 BC = ca. 6,000–5,900 BP). Numerous remains: einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (rare); six-rowed barley (frequent); naked barley (frequent); lentil (frequent). 4. Vršnik, Štip, Macedonia (Hopf 1961a). Early Neolithic (Starčevo group, 4,900 BC = ca. 6,900 BP). Numerous remains: einkorn wheat (prevailing); emmer wheat (frequent); free-threshing wheat (rare); barley (rare). 5. Pokrovnik, Dalmatia (Karg and Müller 1990). Middle Neolithic, Danilo culture (6,290±65 BP = ca. 7,300–7,200 cal BP). Numerous remains in a storage context: einkorn wheat (few); emmer wheat (prevailing). 6. Gomolava, Serbia (van Zeist 1975, 2003). Late Neolithic, Vinča culture (ca. 5,715±75 uncal BP = ca. 5,900–5,700 cal BP). Rich remains: einkorn wheat (prevailing); emmer wheat (frequent); six-rowed barley (frequent); broomcorn millet (frequent); lentil (few); pea (rare); Vicia sp. (rare); flax (rare). Wild: Malus sp.; Cornus mas; Vitis vinifera; Physalis alkekengi (all few); Fragaria vesca (rare); Avena sp. (rare).

Former Yugoslavia (General references: Renfrew 1979; Kroll 1991) 1. Starčevo, Serbia (Renfrew 1979, Table 5). Early Neolithic (ca. 5,000 BC = ca. 7,000 BP). Numerous remains, all from impression: einkorn wheat (few); emmer wheat (prevailing); naked wheat (few); sixrowed barley (few); pea (few). Wild: Malus sp. and Cornus mas (few). 2. Anza, Macedonia (Renfrew 1976). (i) Early Neolithic (7,270±140 to 6,070±200 uncal BP = ca.

Bulgaria (General references: Renfrew 1979; Kroll 1991; Popova 1995; Marinova 2007). 1. Karanovo, Sliven (Thanheiser 1997; Marinova 2004, 2006). (i) Early Neolithic, Phase I–II/III (ca. 8,000–7,550 cal BP). Few remains: einkorn wheat (frequent); emmer wheat (prevailing); free-threshing wheats (few); hulled two/six-rowed barley (frequent); lentil (few); bitter vetch (frequent); grass pea

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(few). Possible weeds: additional ca. 18 taxa. (ii) Middle Neolithic, Phase III (6,510±60 to 6,130±60 uncal BP = ca. 7,500–6,950 cal BP). Rich remains: einkorn wheat (few); emmer wheat (prevailing); free-threshing wheat (few); naked six-rowed barley (storages); grass pea (storages); bitter vetch (storages); lentil (storages). Possible weeds: additional ca. two taxa. (iii) Late Neolithic, Phase III/IV and IV (ca. 6,850–6,400 cal BP). Rich charred remains: einkorn wheat (few); emmer wheat (prevailing); free-threshing wheats (few); naked six-rowed barley (storages); grass pea (storages); bitter vetch (storages); lentil (storages). Possible weeds: additional ca. thirty taxa. (iv) Early Eneolithic, Phase V. Rich remains: einkorn wheat (storages); emmer wheat (few); naked six-rowed barley (prevailing); lentil (rare); bitter vetch (storages). Possible weeds: additional ca. thirty-five taxa. (v) Late Eneolithic, Phase VI (ca. 6,650 cal BP).Rich remains: einkorn wheat (frequent); emmer wheat (frequent); naked six-rowed barley (few); bitter vetch (storages). (vi) Early Bronze Age, Phase VII (ca. 4,650–4,450 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (few); hulled six-rowed barley (prevailing); lentil (rare); bitter vetch (storages); grass pea (rare); safflower (few).Weeds: Galium sp.; Polygonum convolvulus. Wild fruits have been found in almost all phases: Cornus mas (frequent); Vitis vinifera subsp. sylvestris; Prunus avium/cerasus; Rubus ulmifolius/idaeus; Sambucus nigra; Ficus carica; Corylus avellana; Fragaria vesca; Quercus sp. (acorns); Physalis sp. 2. Kovacevo, Blagoevgrad (Popova 1992; Marinova 2006). (i) Early Neolithic, Kovacevo Ia-Id (7,245±36 to 7,028±35 uncal BP = ca. 8,150– 7,850 cal BP). Rich charred remains: einkorn wheat (frequent); emmer wheat (frequent); hulled barley (very frequent); lentil (frequent); grass pea (very frequent); pea (few); bitter vetch (few); chickpea (few). Wild: Cornus mas (very frequent); Vitis vinifera subsp. sylvestris (frequent); Corylus avellana; Prunus avium/cerasus (frequent); Sambucus nigra; Rubus ulmifolius/idaeus (frequent); Pistacia terebinthus. Possible weeds: additional ca. twenty-seven taxa. (ii) Bronze Age. Rich charred remains: einkorn wheat (prevailing); barley (frequent); millet (rare); grass pea (few). Weed: Galium sp.

3. Azmaška (Tell Azmak), Sliven (Hopf 1973a; Renfrew 1979, Table 7). (i) Early Neolithic, Phase I (ca. 7,850–7,500 cal BP). Rich remains: einkorn wheat (few); emmer wheat (prevailing); free-threshing wheat (frequent); naked six-rowed barley (rare); lentil (frequent); pea (few); grass pea (few). (ii) Middle Neolithic, Phase II (6,476±100 uncal BP = ca. 7,450–7,300 cal BP). Rich remains: bitter vetch (pure storage). Wild: Sambucus sp. (iii) Eneolithic (ca. 6,400–6,300 cal. BP). Rich remains: einkorn wheat (frequent); emmer wheat (frequent); naked sixrowed barley (frequent); hulled six-rowed barley (rare); pea (few); lentil (frequent); bitter vetch (prevailing). Wild: Vicia sp. 4. Kapitan Dimitrievo, Pazardzhik (Marinova 2006, forthcoming). (i) Early Neolithic I (ca. 7,950– 7,650 cal BP). Rich charred remains: einkorn wheat (storages); emmer wheat (storages); grass pea (storages). (ii) Early Neolithic II (ca. 7,850–7,700 cal BP). Rich charred remains: einkorn wheat (prevailing, storages); emmer wheat (frequent, storages); hulled barley (storages); naked barley (storages); lentil (frequent); grass pea (storages); bitter vetch (rare); pea (rare); chickpea (rare). Wild: Cornus mas (very frequent); Vitis vinifera subsp. sylvestris; Pistacia terebinthus; Corylus avellana; Prunus avium/cerasus; Sambucus nigra (frequent); Rubus ulmifolius/idaeus (frequent); Physalis sp. Possible weeds: additional ca. thirty-six taxa. (iii) Late Neolithic: einkorn wheat (prevailing); emmer wheat (frequent); naked barley (rare); lentil (frequent); pea (few); bitter vetch (rare); grass pea (prevailing); coriander (few). Wild: Cornus mas (frequent); Corylus avellana; Prunus avium/cerasus (frequent); Sambucus nigra (frequent); Rubus ulmifolius/idaeus (frequent); Physalis sp.; Fragaria vesca. Possible weeds: additional ca. two taxa. (iv) Early Eneolithic. Rich charred remains: einkorn wheat (prevailing); emmer wheat (few); naked wheat (rare); bitter vetch (rare); grass pea (few). 5. Galabovo, south Bulgaria (Popova 1995b; Popova 2001). (i) Chalcolithic culture. Rich remains: einkorn wheat (very frequent); emmer wheat (frequent); spelt wheat (few); free-threshing compactumtype wheat (rare); hulled and naked barley (frequent); bitter vetch (very frequent). (ii) Early Bronze Age culture. Rich remains: einkorn wheat (prevailing); emmer wheat (very frequent); spelt wheat (rare); free-threshing compactum-type wheat

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

(rare); hulled and naked barley (frequent); lentil (very frequent, storages); bitter vetch (very frequent, storages). Weeds: Agrostemma githago; Chenopodium album; Polygonum lapatifolium; Rumex sp. 6. Yunatzite, Pazardzhik (Popova and Pavlova 1994; Marinova and Popova 2008). (i) Late Chalcolithic. Rich charred remains: einkorn wheat (storages); emmer wheat (frequent); free-threshing wheat (rare); naked barley (storages); millet (few); lentil (frequent); bitter vetch (very frequent); grass pea (storages); chickpea (few). Weeds: Agrostemma githago; Chenopodium album; Polygonum convolvulus; Rumex sp. (ii) Bronze Age. Rich charred remains: einkorn wheat (very frequent, storages); emmer wheat (frequent); hulled barley (prevailing, storages); lentil (frequent); bitter vetch (storages); grass pea (storages). Wild: Quercus sp. acorns (frequent); Vitis vinifera subsp. Sylvestris. Weeds: Agrostemma githago; Bromus sp.; Centaurea sp.; Polygonum aviculare; Brassicaceae.

Rumania (General references: Wasylikowa et al. 1991; Cârciumaru 1996; Monah 2007) 1. Liubcova, Caraş Severin district (Cârciumaru 1996). Neolithic Vinča culture B2 (second half of the eighth millennium BP). Rich charred remains: einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (frequent); hulled and naked barley (few); lentil (prevailing). Wild: Galium spurium; Rumex acetosa; Vicia sp. 2. Cârcea, Dolj district (Cârciumaru 1996). Neolithic Dudeşti culture (ca. 6,500–6,000 BP). Rich charred remains: einkorn wheat (few); emmer wheat (few); free-threshing wheat (frequent); barley (few); pea (pure hoard); rye (few). 3. Poduri, Bacău district (Cârciumaru and Monah 1985; Monah and Monah 2008). (i) Late Neolithic, Precucuteni culture (ca. 6,950–6,600 cal BP). Vessels and adobe silos containing large amount (tens of kilograms) of charred grains: einkorn wheat (frequent); emmer wheat (frequent); durum wheat? (rare); spelt wheat (rare); bread wheat (frequent); club wheat (rare); domestic barley (frequent); Coriander (rare). Weeds: Setaria viridis; Vicia cracca; Polygonum convolvulus; Rumex acetosa; Galium sp.; Chenopodium album. Wild:

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Corylus avellana; Tilia platyphyllos. (ii) Eneolithic Cucuteni A culture (ca. 6,650–6,350 cal BP). Einkorn wheat (frequent); emmer wheat (frequent); club wheat (rare); bread wheat (frequent); domestic barley (frequent); oat (rare); rye (rare); pea (rare); grapevine (rare); Prunus domestica (rare); coriander (rare). Weeds: Setaria pumila; Agrostemma githago; Chenopodium album; Rumex acetosa; Brassica nigra; Thlaspi arvense; Galium sp. Wild: Cerasus avium; Rubus idaeus; Malus/Pyrus; Corylus avellana; Rosa sp.; Cornus mas; Sambucus nigra. (iii) Eneolithic Cucuteni B culture (sixth millennium BP). Einkorn wheat (frequent); emmer wheat (frequent); domestic barley (frequent); rye (rare); coriander (rare). Wild: Sambucus nigra; Viburnum lantana; Cornus mas; Rubus sp.; Vicia sativa. 4. Celei (Sucidava), Olt district (Cârciumaru 1996). Transition from Eneolithic to Bronze Age (ca. 4,200 BP). Large quantities of charred grains: einkorn wheat (few); emmer wheat (frequent); spelt wheat (frequent); free-threshing wheat (frequent); six-rowed hulled barley (frequent); lentil (few); lumps of linseed. Wild: Rumex acetosa; Carex rostrata. 5. Tell Hârşova, Constanţa district (Cârciumaru 1996; Monah 2002; Monah and Monah 2008). (i) Late Neolithic Boian culture (seventh millennium BP). Einkorn wheat (frequent); wild emmer wheat? (frequent?); emmer wheat (frequent); spelt wheat (frequent); pea (frequent). (ii) Late Neolithic/ Eneolithic Gumelniţa Culture (ca. 6,350–6,150 cal BP). Einkorn wheat (few); emmer wheat (few); bread wheat (few); domestic barley (few); rye (few); lentil (frequent); pea (frequent); bitter vetch (prevailing); grapevine (rare). Wild: Vicia sp.; Sambucus nigra; Vitis sylvestris.

Moldavia and Ukraine (General references: Januševič 1984; Wasylikowa et al. 1991; Pashkevich 2003) 1. Sacarovca, Sângerei district, Moldavia (Januševič 1984; Kuzminova et al. 1998). Early Neolithic, Starčevo-Criş culture (6,650±50 uncal. BP = ca. 7,600–7,500 cal BP). Numerous remains, both imprints and charred grains: einkorn wheat (frequent); emmer wheat (prevailing); spelt wheat (few); hard wheat, T. aestivum/compactum (rare); oat

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DOMESTICATION OF PLANTS IN THE OLD WORLD

(rare); domestic barley (frequent); broomcorn millet (frequent); pea (few); cherry plum (rare); plum (rare). Weeds: Alyssum sp.; Agrostemma sp.; Lathyrus sp.; Setaria viridis; Setaria glauca; Galium sp. Wild: Coryllus avelana; Quercus robur; Cornus mas; Malus sp.; Vitis sylvestris. 2. Ruseştii Noi (Novye Rusešty), Chişinău district, Moldavia (Januševič 1976, 1986).(i) Linearbandkeramik (ca. 7500 BP), ear impressions in pottery: emmer wheat; spelt wheat. (ii) Precucuteni (Tripolye A, early phase) culture (5,615±100 uncal BP = 6,500–6,300 cal BP): impressions and charred grains: einkorn wheat; emmer wheat; spelt wheat; hard wheat; naked barley. (iii) Cucuteni A (Tripolye B, middle phase), impressions and charred grains: emmer wheat; spelt wheat; naked barley; grapevine. 3. Putineşti, Soroca district, Moldavia (Januševič 1976). Eneolithic, Precucuteni (ca. 6,000 BP). Impressions and charred grains: einkorn wheat; emmer wheat; spelt wheat; hard wheat; barley; naked barley; oat; broomcorn millet? Wild: Cornus mas; Pyrus elaegrifolia; Cerasus avium. 4. Cuconeştii Vechi (Starye Kukoneshti), Moldavia (Januševič 1978, 1984; Kuzminova 1988). Eneolithic, Cucuteni A–B (Tripolye culture, middle phase). Rich remains (mainly carbonized, few imprints): einkorn wheat (few); emmer wheat (prevailing); naked barley (few). Weeds: Rumex acetosella; R. acetosa; Chenopodium album; Saponaria officinalis; Sinapis arvensis; Polygonum aviculare; Echinochloa crus-galli; Medicago falcata; Solanum nigrum; Stellaria media; Bromus secalinus; Datura stramonium; Glaucium corniculatus. 5. Rivne (Rovno), Rivne Oblast, Ukraine (Pashkevich 2003). Linearbandkeramik (ca. 6,500–6,000 BP). Rare imprints: emmer wheat; bread wheat; naked barley; broomcorn millet; pea. 6. Eneolithic settlements in Ukraine, Tripolye cultures (ca. 6,000–4,750 BP). Luca Vrublevecaja (Januševič 1976), Majdaneckoe, Cherkas’ka Oblast, Majaki and Usatovo, Odessa Oblast (Pashkevich 2005). Few remains, mostly imprints, some charred: einkorn wheat; emmer wheat; spelt wheat; bread wheat; hard wheat; naked barley; broomcorn millet; pea; plum; cherry plum; apricot. Wild: Vitis sp.; Pyrus/Malus; Corylus avellana.

Hungary (General references: Tempír 1964; Hartyáni and Nováki 1975; Füzes 1990; Wasylikowa et al. 1991; Gyulai 2007). 1. Körös-Starčevo culture sites (Szeged-Gyálarét, Röszke-Lúdvár, Battonya-Basarága) (Hartyányi et al. 1968; Hartyányi and Nováki 1971; Hartyáni and Nováki 1975; Füzes 1990). Early Neolithic (ca. 7,950–7,250 cal BP). Imprints: einkorn wheat (prevailing); emmer wheat (few). Probable weeds: Lathyrus sp.; Bromus sp. Wild: Cornus mas; Corylus avellana. 2. Füzesabony-Gubakút, Northern Great Plain (Gyulai 2007). Middle Neolithic LBK culture, Great Hungarian Plain Group (ca. 7,550–7,150 cal BP). Scarce charred remains: emmer wheat (rare); barley (frequent). Weed: Avena fatua. 3. Zánka-Vasútállomás (Füzes 1990, 1991). Middle Neolithic LBK culture, Trans-Danubian Group (ca. 7,250–6,650 cal BP). Rich charred remains and imprints: einkorn wheat (including wild forms), grains, and rachis fragments (very common); emmer wheat, grains, and rachis fragments (very common); bread wheat; club wheat (Triticum aestivum subsp. compactum); spelt wheat, spikelets; two-rowed barley grains; common millet; pea. Weeds: Agrostemma githago; Bromus cf. sterilis; Galium sp.; Fallopia convolvulus; Chenopodium sp.; rye brome; T. baeoticum? 4. Dévaványa-Réhelyi dűlő (Hartyányi et al. 1968). Middle Neolithic LBK culture, Eastern region, Szakálhát-Szilmeg Group (ca. 6,350 cal BP). Rich charred remains: einkorn wheat (few); emmer wheat (frequent); barley (prevailing); lentil (few). Weeds: Fallopia convolvulus; Chenopodium sp. 5. Berettyóújfalu-Szilhalom (Hartyányi et al. 1968). Late Neolithic, Tisza-Herpály-Berettyóvölgy site. Rich charred remains: einkorn wheat (frequent); emmer wheat (few); bread wheat (rare); two-rowed naked barley (frequent); common millet; field pea (frequent); bitter vetch (rare). Wild: Malus sylvestris fruits. 6. Rákoskeresztúr-Újmajor (Gyulai 1999). Eneolithic (Middle Copper Age), Ludanice culture (ca. 6,250–5,450 cal BP). Rich remains: emmer wheat (rare); six-rowed barley (few). Wild: Rumex conglomeratus; Schoenoplectus lacustris; Thalicrum flavum; Chenopodium sp.

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

7. Budapest. (i) Csepel-Vízmű (Endrődi and Gyulai 2000). Eneolithic (Late Copper Age), Baden culture (ca. 5,450–4,950 cal BP). Rich charred remains: einkorn wheat (few); emmer wheat (rare); club wheat (rare); barley (rare). Weeds: Bromus arvensis; B. secalinus; Chenopodium sp.; Fallopia convolvulus. Wild: Crataegus monogyna; Fagus silvatica; Quercus robur/petraea. (ii) Albertfalva (Gyulai 2003). Early Bronze Age (ca. 4,950–3,550 cal BP), Bell Beaker culture Csepel Group. Rich charred remains: einkorn wheat (frequent); emmer wheat (frequent); bread wheat (rare); spelt wheat (rare); six-rowed naked barley (few); two-rowed barley (frequent); pea (rare); small seed fava bean (rare). Wild: crab apple; many weeds. 8. Bölcske-Vörösgyír (Berzsényi and Gyulai 1998). Middle Bronze Age, Nagyrév and Vatya culture (ca. 3,550–3,250 cal BP). Rich charred remains: einkorn wheat (prevailing); emmer wheat (frequent); spelt wheat (few); bread wheat (rare); sixrowed barley (frequent); two-rowed barley (frequent); naked barley (few); common millet (few); small seed lentil (frequent); bitter vetch (frequent); pea (few); small seed fava bean (few). Weeds: Avena fatua; Bromus arvensis; B. mollis; B. secalinus; B. tectorum; Chenopodium sp.; Fallopia convolvulus; Setaria viridis. Wild: Cornus mas; Crataegus monogyna; Prunus spinosa; Quercus sp.

Austria (General references: Küster 1991; Kohler-Schneider 2007) 1. Schletz near Asparn, Lower Austria (Schneider 1994; Kohler-Schneider 2007). Late Linearbandkeramik culture (6,175±65–6,025±55 uncal BP=ca. 7,150– 6,800 cal BP). Major fortified site of the Early Neolithic. Rich charred remains: einkorn wheat, mainly twograined (very frequent); emmer wheat (very frequent); naked wheat (few); hulled barley (few); pea (few); lentil (rare); flax (linseed); cf. Camelina sativa; Papaver somniferum spp. setigerum. Wild: Cornus mas; Sambucus nigra; Sambucus ebulus; Solanum dulcamara; Physalis alkekengi. Numerous weeds and herbs: Chenopodium album; Ch. hybridum; Fallopia convolvulus; Polygonum aviculare; Bromus secalinus; wild Paniceae. 2. Mondsee, lake-shore settlements, east of Salzburg, Upper Austria (Hofmann 1924; Kohler-

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Schneider 2007). Late Neolithic (4,910±130–4,750±90 uncal BP=ca. 5,750–5,450 cal BP). Rich charred and waterlogged remains: einkorn wheat (few); emmer wheat (very frequent); hulled six-rowed barley (very frequent); pea (rare); flax (few); poppy (few). Wild: Corylus avellana;Malus sylvestris (entire and halves fruits); Rosa canina; Fagus sylvatica; Quercus robur; Tilia platyphyllos; Rubus idaeus; R. fruticosus; Fragaria vesca; Daucus carota; Pastinaca sativa. 3. Anzingerberg/Hundssteig, near Krems, Lower Austria (Kohler-Schneider 2007; KohlerSchneider and Caneppele 2009). Late Neolithic Jevišovice culture (ca. 5,050–4,750 cal BP). Rich charred remains: einkorn wheat; emmer wheat; spelt wheat (few, very early); free-threshing wheat (rare); hulled barley; broomcorn millet (frequent); foxtail millet (few); lentil (rare); pea (rare); flax (rare); poppy (very frequant). Wild: Vitis vinifera subsp. sylvestris; Prunus spinosa; Rubus fruticosus; R.; Fragaria vesca; Viburnum opulus; Camelina microcarpa; Daucus carota; Atropa belladonna; Hyoscyamus niger. Additional ca. 100 wild taxa from various habitats. 4. Stillfried, Lower Austria (Kohler-Schneider 2001; Kohler-Schneider 2003). Late Bronze Age hillfort site (ca. 3,200–2,700 cal BP). Rich charred remains from storage pits and other domestic contexts: einkorn wheat (very frequent); emmer wheat (frequent); spelt wheat (very frequent); free-threshing wheat (rare); hulled barley (very frequent); rye (few); broomcorn millet (very frequent); foxtail millet (frequent); lentil (frequent); pea (rare); fava bean (few); bitter vetch (rare); gold of pleasure (frequent); poppy (rare); grapevine (few). Wild: Prunus avium/ cerasus; Malus/Pyrus; Fragaria sp.; Digitaria sanguinalis; Daucus carota; Physalis alkekengi. Additional ca. 100 wild taxa from various habitats. 5. Dürrnberg/Hallein, Salzburg (Werneck 1949; Swidrak 1999; Boenke 2007). Iron Age salt mining site and settlement (ca. 2,400–2,050 cal BP). Rich remains from human feces (salt preservation), charred and waterlogged material: emmer wheat (few); spelt wheat (few); free-threshing wheat (rare), hulled and naked barley (frequent); broomcorn millet (very frequent); foxtail millet (few); lentil (few); pea (few); broad bean (few); poppy (frequent); gold of pleasure (few); hemp, fruits, and fibres (frequent); flax, seeds, and fibres (few). Wild: Corylus avellana; Prunus domestica subsp. insititia; P. spinosa; P. avium;

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Malus sylvestris; Pyrus sp.; Rubus fruticosus; R. caesius; R. idaeus; Fragaria vesca; Rosa sp.; Crataegus laevigata;Vitis vinifera cf. subsp. sylvestris; Sambucus nigra; Viburnum lantana; Daucus carota; Carum carvi.

Italy (General references: Rottoli and Pessina 2007; Rottoli and Castiglioni 2009) 1. Grotta dell’Uzzo, Sicily (Costantini, 1989). (i) Mesolithic (ca. 9,950–8,950 uncal BP = ca. 11,400– 10,150 cal BP). Scarce remains, all wild: Arbutus unedo; legume seeds (probably Lathyrus or Pisum); Quercus acorns; two pips of Vitis. (ii) Early Neolithic (6,750±70 uncal BP = ca. 7,650–7,550 cal BP). Scarce remains (unspecified quantities): einkorn wheat; emmer wheat; lentil; barley (somewhat later). (iii) Middle Neolithic (ca. 6,750–6,450 uncal BP = ca. 7,600–7,350 cal BP). Few remains: einkorn wheat; emmer wheat (prevailing); free-threshing wheat; barley (relatively common); lentil; pea; bitter vetch; faba-like bean. Wild: Amygdalus sp; Olea europaea; Vitis vinifera; Ficus carica. 2. Scamuso, Torre a Mare, Bari (Costantini et al. 1997). Numerous remains including chaff: (i) Impressed Ware culture (ca. 7,150–6,750 uncal BP = ca. 8,000–7,600 cal BP): einkorn wheat (few); emmer wheat (frequent); hulled barley (prevailing); lentil (few); pea (few). Wild: Vitis vinifera (few); Lathyrus sp. (frequent). (ii) Painted Ware culture (ca. 6,600– 5,800 uncal BP = ca. 7,500–6,550 cal BP): einkorn wheat (rare); emmer wheat (frequent); free-threshing wheat (few); barley (prevailing); lentil (rare); pea (rare); faba-like bean (a single seed). Wild: Avena sp. (rare); Vitis vinifera (few); Lathyrus sp. (few); few herbs. 3. Pienza, Siena (Castelletti 1976). (i) Impressed Ware (Cardial) culture (ca. 7,750–7,250 cal BP). Scarce remains: einkorn wheat (rare); emmer wheat (prevailing); free-threshing wheat (frequent); hulled/naked barley (few). Wild: Avena sp.; Vitis sylvestris. (ii) Middle Neolithic (ca. 6,950–6,450 uncal. BP = ca. 7,800–7,350 cal BP). Scarce remains: emmer wheat (rare); free-threshing wheat (rare); barley (rare). Wild: cf. Avena. (iii) Late Neolithic (ca. 6,450–5,950 uncal. BP = ca. 7,400–6,750 cal BP). Scarce remains: emmer wheat (rare); free-threshing wheat (rare), barley (rare). Wild: Avena sp. (iv)

Bronze Age: Scarce remains: einkorn wheat (few); emmer wheat (prevailing); hulled barley (few); broomcorn millet (few); faba bean (few). Wild: Avena sp.; Cornus mas. 4. Sammardenchia, and other adjacent Early Neolithic sites in Friuli, northern Italy (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007). Early north Italian Neolithic (6,570±74 uncal BP to 5,684±58 uncal BP= ca. 7,550–6,450 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (rare); six-rowed hulled barley (frequent); lentil (frequent); pea (frequent); vetches (Vicia sativa agg.) (frequent); bitter vetch (few); grass pea (few). Rye, spelt, and millets apparently infested grain agriculture. Wild: Vitis vinifera; Corylus avellana; Cornus mas; Malus sylvestris; Quercus sp.; Rubus fruticosus agg.; Crataegus sp.; Prunus insititia/spinosa; Juglans regia; Bromus spp. (frequent). Weeds: Galium aparine; Rumex sp.; Fallopia convolvulus. 5. San Marco, Gubbio, Perugia (Costantini and Stancanelli 1994). Early-Middle Neolithic (6,300– 5,750 uncal BP = ca. 7,250–6,500 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (prevailing); compactum-type wheat (frequent); hulled barley (few); naked barley (rare); pea (rare). Wild: Corylus avellana (frequent); Ficus carica; Prunus insititia; Vitis vinifera. 6. Monte Còvolo, Villa Nuova sul Clisi, Brescia (Pals and Voorrips 1979). (i) Late Neolithic (ca. 5,850–5,450 uncal BP = ca. 6,700–6,200 cal BP). Scarce remains: einkorn wheat (rare); emmer wheat (prevailing); six-rowed barley (few). Wild: Cornus mas; Malus sylvestris; Prunus avium; P. spinosa; Vitis sylvestris; Quercus sp.; Rosa sp.; Rubus sp.; Physalis alkekengi. (ii) White Ware phase (ca. 5,500–4,150 uncal BP = ca. 6,300–4,600 cal BP). Scarce remains: einkorn wheat (few); emmer whet (prevailing); sixrowed barley (frequent); broomcorn millet (rare). Wild: Prunus avium; Quercus sp.; Vitis sylvestris; Rosa sp; Rubus sp.; Physalis alkekengi. (iii) Bell Beaker phase (ca. 4,150–3,900 uncal BP = ca. 4,800–4,300 cal BP). Numerous remains: einkorn wheat (few); emmer wheat (prevailing); six-rowed barley (frequent). Wild: Cornus mas; Malus sylvestris; Prunus avium; P. spinosa; Quercus sp.; Vitis sylvestris; Physalis alkekengi. (iv) Early Bronze Age (ca. 3,900–3,350

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

uncal BP = ca. 4,400–3,550 cal BP). Scarce remains: emmer wheat (prevailing); six-rowed barley (few). Wild: Malus sylvestris. 7. La Marmotta, Lake Bracciano, Rome (Rottoli 1993, 2002). Rich Early Neolithic (6,855±65 uncal BP to 6,310±75 uncal BP = ca. 7,750–7,150 cal BP) waterlogged remains: einkorn wheat; emmer wheat; freethreshing turgidum-type wheat; hulled two-rowed barley; lentil; pea; grass pea; vetches; flax (rare). Wild: Vitis vinifera (frequent); Papaver somniferum subsp. setigerum (very frequent); Corylus avellana; Malus sylvestris.; Pyrus sp.; Quercus sp.; Ficus carica; Prunus spinosa/domestica; Fragaria vesca; Cornus mas; Carthamus lanatus; Silybum marianum; Crataegus sp.; Rubus fruticosus agg.; Cornus sanguinea; Laurus nobilis.

Poland (General references: Wasylikowa et al. 1991; Bieniek 2007; Lityńska-Zając 2007) 1. Strachów, Silesia (Lityńska-Zając 1997). (i) Linearbandkeramik culture (6,535–6,170 uncal BP = ca. 7,350–7,050 cal BP). Numerous remains (imprints and some charred grains): einkorn wheat (few); emmer wheat (prevailing); barley (few). Wild: various grasses and herbs such as Bromus sp.; Setaria viridis/verticillata; Echinochloa crus-galli; Polygonum sp.; Rumex sp.; Chenopodium sp.; Bromus cf. tectorum; Chenopodium album; Polygonum convolvulus; Rumex sp. (ii) Lengyel-Polgár culture (ca. 5,730 uncal BP = ca. 6,450 cal BP). Scarce remains (charred and imprints): einkorn wheat (few); emmer wheat (few); barley (very few). (iii) Funnel Beaker culture (ca. 4,800–4,480 uncal BP = ca. 5,550–5,050 cal BP). Some remains, mostly imprints: einkorn wheat (rare); emmer wheat (prevailing); barley (rare). Wild: various grasses and herbs; among others, Bromus ssp. and Chenopodium album. 2. Nowa Huta-Mogila, Kraków district (Gluza 1984). Neolithic, Lengyel culture (ca. 5,430 uncal BP = ca. 6,300–6,200 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (prevailing); naked six-rowed barley (frequent). Large variety of herbaceous weedy species: Bromus arvensis (very aboundant); Bromus racemosus (abundant); several species of Chenopodium, Polygonum and Galium; Fallopia convolvulus (abundant); Digitaria ssp.; Echinochloaa crusgalli; Setaria viridis/verticillata; Solanum nigrum;

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Trifolium ssp.; Wild: Corylus avellana; Sambucus ebulus. 3. Gniechowice and Stary Zamek (Gluza 1994). Linearbandkeramik culture (ca. 6,950 uncal BP = 7,800–7,750 cal BP). Mostly imprints: einkorn wheat (few); emmer wheat (prevailing); barley (few); Bromus sp. (few). 4. Ćmielów, Tarnobrzeg district (Klichowska 1976). Funnel Beaker culture (ca. 5,600–5,300 cal BP). Rich remains: emmer wheat (prevailing); pea (few); lentil (few); linseed (frequent). Wild: Chenopodium album; Bromus secalinus; Rumex acetosa; Setaria glauca; Berberis vulgaris; Taxus baccata. 5. Gwoździec, Małopolska (Bieniek and LitynskaZajac 2001; Lityńska-Zając 2007). Linear Pottery culture (ca. 7,400–6,600 cal BP). Numerous charred and imprinted remains: einkorn wheat (frequent); emmer wheat (prevailing); naked barley (rare). Wild: Chenopodium album;Bromus sp.; Echinochloa crus-galli; Setaria pumila; Malus sylvestris (pips and fruit fragments). 6. Brześć group, Kujawy (Bieniek 2007). (i) Linearbandkeramik culture (ca. 7,450–6,950 cal BP). Numerous charred remains: einkorn wheat, mainly chaff (prevailing); emmer wheat, mainly chaff (frequent); hulled barley (rare); flax (rare); poppy (very rare). Wild: Chenopodium album; Fallopia convolvulus; Echinochloa crus-galli; several species of Polygonum; Physalis alkegengi; Solanum nigrum;Schoenoplectus tabernaemonatani; and other species. (ii) Lengyel culture (ca. 6,350–5,950 cal BP). Rich charred remains: einkorn wheat (frequent); emmer wheat (prevailing); bread wheat (rare); hulled barley (rare). Wild: several species of Bromus and Galium; Chenopodium album; Fallopia convolvulus; Hierochloe cf. australis; Stipa pennata; and other species. (iii) Funnel Beaker culture (ca. half of the sixth millennium BP). Numerous charred remains: einkorn wheat, mainly chaff (prevailing); emmer wheat, chaff (rare); poppy (rare). Wild: Chenopodium album; Fallopia convolvulus; several species of Polygonum; and others.

Czech Republic and Slovakia (General references: Hajnalová 1989; Wasylikowa et al. 1991; Hajnalová 2007) 1. Mohelnice, Litovel, Moravia (Opravil 1979; Kühn 1981; Opravil 1981). Linearbandkeramik

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DOMESTICATION OF PLANTS IN THE OLD WORLD

culture (6345±100 to 622±80 uncal BP = ca. 7,400– 7,000 cal BP). Numerous remains: einkorn wheat (rare); emmer wheat (prevailing); free-threshing wheat (frequent); two-rowed barley (few); broomcorn millet (rare); pea or vetch (rare); flax cord (few). Wild: Malus sylvestris; Corylus avellana. 2. Blatné near Štrky, Bratislava district, Slovakia (Hajnalová 1989). (i) Linearbandkeramik culture (ca. 7,250–7,000 cal BP). Numerous remains: einkorn wheat (frequent); emmer wheat (prevailing); spelt wheat (few); pea (rare). (ii) Middle Neolithic, Zeliezovce group (ca. 6,950–6,650 cal BP). Numerous remains: einkorn wheat (frequent); emmer wheat (prevailing); spelt wheat (few); hulled six-rowed barley (rare); pea (rare); lentil (few). 3. Bylany, Kutná Hora, Bohemia (Tempír 1979). (i) Linearbandkeramik culture (ca. 7,400–7,000 cal BP), Stroked pottery culture (ca. 7,000–6,550 cal BP). Numerous remains: einkorn wheat (few); emmer wheat (prevailing); broomcorn millet (few); pea (few); lentil (few). (ii) Funnel Beaker culture (ca. 6,950–5,150 cal BP). Rich remains: einkorn wheat (prevailing); emmer wheat (few); free-threshing wheat (few); six-rowed hulled barley (frequent). Wild: several weeds. 4. Šarišské Michalˇany-Fedelemka, Sabinov, Prešov, Slovakia (Hajnalová 1993; Hajnalová and Hajnalová 2004). Middle Neolithic (Bükk culture, ca. 6,950–6,650 cal BP). Numerous charred remains: einkorn wheat (frequent); emmer wheat (frequent); spelt wheat (few); free-threshing wheat (few); hulled and naked barley (frequent); rye (few); broomcorn millet (few); pea (few); lentil (few); flax (few). Wild: Malus sp.; cf. Cerasus avium; Prunus sp.; Rubus sp.; Trapa natans. 5. Bajč-Medzi kanálmi, Komárno, Nitra, Slovakia (Cheben and Hajnalová 1997). Middle Neolithic (Želiezovce group, ca. 6,950–4,700 cal BP). Rich charred and imprints, mostly from in situ content of an oven: one- and two-seeded einkorn (prevailing); emmer wheat (few); spelt wheat (few); free-threshing wheat (rare); naked six-row barley (rare). Weed seeds. 6. Nitriansky Hrádok, Nové Zámky, Slovakia (Tempír 1969; Kühn 1981). (i) Early Bronze Age (Maďarovce culture, ca. 1,950–1,400 cal BC = ca. 3,900–3,350 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (prevailing); free-thresh-

ing wheat (frequent); six-rowed and two-rowed barley (frequent); oat (rare); rye (rare); pea (few/ pure); lentil (rare). Wild: Avena fatua; Bromus secalinus; Lolium sp.; Vicia sp.; Agrostemma githago; Quercus robur. (ii) Bronze Age. Numerous remains: einkorn wheat (rare); emmer wheat (rare); six-rowed barley (prevailing); pea? Wild: Bromus secalinus; B. arvensis; Lolium temulentum; Agrostemma githago; various Cruciferae.

Switzerland (General references: Jacomet 2006, 2007, 2008; Jacomet and Brombacher 2005; Jacomet and Behre 2009). 1. Egolzwil 3, Wauwiler Moos, Canton of Lucerne (Bollinger 1994; Jacomet 2007). Early Late Neolithic Egolzwil culture (ca. 5,650 uncal BP = ca. 6,250 cal BP). Lake-shore settlement at a small shallow lake. Rich remains, mostly waterlogged: einkorn wheat (rare); emmer wheat (rare); tetraploid free-threshing wheat (frequent); six-rowed barley (frequent); pea (frequent); poppy seeds (very frequent); flax, capsules and seeds (few). Wild: hazelnut; crab apple; blackberry and raspberry; wild strawberry (Fragaria vesca); rose hips; Brassica rapa (frequent); additional some 100 taxa—the majority could have been of use. 2. Zürich (town and lake), numerous lake-shore settlements (Jacomet 1988, 2004; Jacomet et al. 1989; Brombacher and Jacomet 1997; Favre 2002; Brombacher et al. 2005). Very rich (several hundreds of thousands) charred or excellently preserved waterlogged plant remains, from several sites covering the various phases of the lake-shore agricultural settlements. (i) Late Neolithic cultures Egolzwil, Early to classical Cortaillod, Pfyn, Horgen, Schnurkeramik (Corded ware) (ca. 6,250–4,450 cal BP, mostly dendrodates). Einkorn wheat (rare); emmer wheat (rare in the earlier phases, prevailing in the later phases); durum/turgidum-type tetraploid wheat (prevailing in the earlier phases, frequent in the later phases); six-rowed, mostly naked barley (frequent); pea (rare, present from Egolzwil culture (ca. 6,350 cal BP) onward; flax, seeds, and capsules (becomes frequent from ca. 5,750 cal BP onwards); poppy (frequent until ca. 4,750 cal BP, then becomes rare). Mediterranean aromatic plants (from ca. 5,900

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

cal BP onwards): celery Apium graveolens (rare); dill Anethum graveolens (rare); lemon balm Melissa officinalis (rare). Large quantities of many plants collected from the wild, particularly: Malus sylvestris; Corylus avellana; Prunus spinosa; Rubus idaeus; R. fruticosus; Fragaria vesca; Brassica rapa; as well as remains of numerous weeds and local herbs. (ii) Bronze Age sites (Mozartstrasse, Alpenquai, and Wädenswil-Vorder-Au). Different phases of the Early Bronze Age from ca. 3,850 cal BP onwards, and Late Bronze Age, ca. 3,000–2,800 cal BP, dendrodates). Same crop species as in the Late Neolithic sites, with the addition of spelt wheat (Triticum spelta) from Early Bronze Age onwards, and broomcorn millet (Panicum milieaceum), foxtail millet (Setaria italica), and lentil (Lens culinaris). At Alpenquai also Camelina was frequent. 3. Toos-Waldi, Canton of Thurgau (Jacomet and Behre 2009). Dryland site on a hilltop, Middle Bronze Age. Rich charred remains: einkorn wheat (rare); emmer wheat (frequent); naked tetra/hexaploid wheat (rare); spelt wheat (frequent); six-rowed hulled barley; whole stocks (frequent). Wild: Avena sp.; Quercus sp., many acorn halves. 4. Canton of Grisons (Graubünden), south-east Switzerland (Jacomet et al. 1998, 1999). Rich plant remains from Early Bronze Age onwards from six sites, the most important one is Savognin-Padnal. (i) Early to Middle Bronze Age (ca. 4,150–3,200 cal BP). Einkorn wheat (few); emmer wheat (frequent); spelt wheat (very frequent at one site); hulled sixrowed barley (prevailing); pea (frequent); faba bean (many at one site); flax (few). (ii) Late Bronze Age (ca. 3,200–2,700 cal BP): einkorn wheat (rare); spelt wheat (rare); hulled six-rowed barley (prevailing); broomcorn millet (rare); pea (few). Wild: Corylus avellana; Quercus sp.; Rosa sp.; Avena fatua; and seeds of other weeds. 5. Lake Biel, Lake-shore settlements, particularly Twann, Nidau-BKW, and Port-Stüdeli, Sutz, Lattrigen, and Lüscherz (Ammann et al. 1981; Brombacher 1997, 2000; Brombacher and Jacomet 2003). (i) Late Neolithic, different phases of the Cortaillod culture (ca. 5,800–5,550 BP, dendro-dates). Rich waterlogged and charred remains, often pure hoards: einkorn wheat (rare to frequent, also two-grained cultivars present); emmer wheat (rare); tetraploid free-threshing wheat (often prevailing); six-rowed barley, mostly

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naked (very frequent); pea, seeds, and pod fragments (in some layers of Twann frequent); flax (frequent, stems in Port-Stüdeli); poppy (frequent); dill (rare, at two sites). Wild: many gathered plants such as Malus sylvestris; Corylus avellana; Rubus idaeus; R. fruticosus; Fragaria vesca; Physalis alkekengi; Brassica rapa; Rosa sp.; Quercus sp. (ii) Different Late Neolithic phases after 5,450 cal BP (Lattrigen, Horgen culture, ca. 5,410–5,176 BP, dendro-dates). Rich waterlogged and charred remains: einkorn wheat (very rare); emmer wheat (prevailing at one site); free-threshing tetraploid wheat (very frequent); six-rowed barley, apparently naked (very frequent); flax (frequent); poppy (frequent); celery and dill (rare, at two sites). Wild: same assemblage as in (i). 6. Lake Neuchatel, shore and surroundings (Villaret-Von Rochow 1971a, 1971b; BaudaisLundström 1978; Schlichtherle 1985; Märkle 2000; Karg and Märkle 2002; Mermod 2007). (i) Late Neolithic to Early Bronze Age (ca. 5,659–4,400 BP, dendrodates). Dryland and lake-shore sites, particularly St Aubin Derrière La Croix, Concise-sousColachoz, Yverdon Avenue des Sports, and St Blaise Bains des Dames. Rich waterlogged and charred plant remains, including stocks: einkorn wheat (few to prevailing); emmer wheat (few to prevailing); tetraploid naked wheat (usually many); six-rowed barley (usually many); celery and dill (rare); pea (rare); flax (frequent); poppy (frequent). (ii) Cortaillod sur les Rochettes Est (Akeret, 2005). Final Neolithic, Bell Beaker culture (ca. 4,250 uncal BP = ca. 4,850 cal BP). Rich remains: einkorn wheat (few); emmer wheat (few); spelt wheat, grains, and chaff (frequent); barley (rare); lentil (rare); bitter vetch; flax. Wild: rich spectrum, including Corylus avellana; Malus sylvestris; Prunus spinosa; Rosa sp.; Rubus sp. 7. Canton of Valais, valley of Valais (Wallis). (i) Sites in Sion (Martin et al. 2008b; Lundström-Baudais in press). Middle Neolithic (ca. 6,450–5,450 cal BP). Few charred remains: einkorn wheat (prevailing); emmer wheat (frequent); naked wheat (few); barley (rare); pea (rare). (ii) Brig-Glis, Waldmatte (Lundström-Baudais in Curdy et al. 1993). Early Iron Age (2550–2450 uncal BP = ca. 2750–2450 cal BP). Rich charred material from a burnt storehouse: barley (frequent); broomcorn millet (frequent); foxtail millet (few); lentil (few); pea (few); bitter vetch (frequent); faba bean (few). No wheat remains.

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Germany (General references: Kreuz 1990; Knörzer 1991; Küster 1991; Kreuz and Boenke 2002; Bogaard 2004; Kreuz et al. 2005; Jacomet 2007, 2009; Bittmann pers. comm.) 1. Hienheim, Kr. Kelheim (Bakels 1978). Linearbandkeramik culture (6,155±45 to 5,910±50 uncal BP = ca. 7,150–6,650 cal BP). Rich remains: einkorn wheat and emmer wheat (prevailing, apparently in equal proportions); pea (frequent); lentil (few); flax (rare). Wild: Corylus avellana; numerous weeds. 2. Heilbronn, Württemberg, including HeilbronnBöckingen, H.-Gross-Gartach, H.-Willsbach (Bertsch and Bertsch 1949); and H.-Klingenberg (Stika 1996a). (i) Linearbandkeramik culture. Rich remains: einkorn wheat (frequent); emmer wheat (frequent); hulled six-rowed barley (frequent); naked six-rowed barley (few); lentil (few); pea (few, except for being very frequent in H.-Klingenberg); flax (rare). (ii) Late Neolithic (ca. 4,750–4,650 uncal BP=ca. 5,600– 5,400 cal BP). Rich remains (both grain and chaff): einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (very frequent); naked barley (very frequent); hulled barley (few); lentil (few); pea (very frequent); poppy (rare); flax (rare). 3. Aldenhovener Platte, Langweiler, Lamersdorf, Bedburg-Garsdorf. Meckenheim and Rödingen (Knörzer 1973, 1974, 1997). Linearbandkeramik culture (5,950±140 to 6,370±210 = ca. 7,500–6,550 cal BP). Rich remains: einkorn wheat and emmer wheat (frequent); pea (few); lentil (rare); flax (few; in one place very frequent); poppy (few). Wild: Bromus secalinus (abundant); Vicia sp.; Quercus sp.; Prunus insititia; P. spinosa; Malus sp.; Corylus avellana; Sambucus sp. 4. Kückhoven, Erkelenz, Rheinland (Knörzer 1998). Linearbandkeramik well (7,040 cal BP, dendrodates). Rich remains, including chaff: einkorn wheat (frequent); emmer wheat (prevailing); freethreshing wheat (few); pea (rare); flax (frequent); poppy (frequent). Wild: Corylus avellana; Malus sylvestris; Prunus spinosa; Rubus caesius; R. fruticosus; Sambucus nigra, and numerous herbs. 5. Dresden-Nickern (Baumann and SchultzeMotel 1968). Linearbandkeramik culture (5,945±100 and 5,815±100 uncal BP = ca. 6,900–6,500 cal BP).

Rich remains: emmer wheat (few); pea (a pure hoard). 6. Dannau, Oldenburg (Kroll 1981b). Middle Neolithic. Rich remains: einkorn wheat (rare); emmer wheat (frequent); naked six-rowed barley (prevailing). Wild: Corylus avellana; Rubus idaeus; Sambucus nigra. 7. Kr. Ludwigsburg sites. (i) BietigheimBissingen (Piening 1989). Early Neolithic, Linearbandkeramik culture (ca. 7,450 cal BP). Rich remains: einkorn (prevailing); emmer wheat (frequent); naked wheat (rare); barley (rare); pea (rare). (ii) Eberdingen-Hochdorf (Küster 1983, 1985; Breuning 1987). Middle Neolithic. Schussenried group (ca. 6,150–ca. 5,750 uncal BP=ca.7,150–6,500 cal BP). Rich remains: einkorn wheat (almost as frequent as naked barley); emmer wheat (few); freethreshing wheat (few); naked six-rowed barley (prevailing); pea (few); linseed (frequent); poppy (frequent). Possibly cultivated: Brassica rapa; Petroselinum crispum. Wild: Corylus avellana; Fragaria vesca; Malus sp.; Sambucus. 8. Federseeried, Württemberg, Lake-shore settlements. (i) Late Neolithic, Aichbühl-Schussenried group (ca. 5,900 cal BP) (Blankenhorn and Hopf 1982). Very rich remains: einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (prevailing); spelt wheat (few); hulled six-rowed barley (few); naked six-rowed barley (rare); poppy (rare). Wild: Corylus avellana; Fagus sylvatica; Malus sp.; Fragaria vesca; Rubus idaeus; R. fruticosus. (ii) Ödenahlen (Maier 1995). Late Neolithic (first half of the fifth millennium BP). Rich remains, both charred and waterlogged, including chaff: einkorn wheat (few); emmer wheat (prevailing); free-threshing wheat (frequent); barley (few); flax (frequent); poppy (rare). Wild: fruits of Malus sp. cut to halves or quarters (numerous); Fragaria vesca; Rubus idaeus. 9. Lake Constancesites (i) Hilzingen (Stika 1991). Early Neolithic, Linearbandkeramik culture (ca. 7,250–7,050 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (frequent); barley (rare); pea (prevailing, in storage); flax (rare). Wild: Corylus avellana; Malus sylvestris; Prunus spinosa. (ii) Hornstaad, lake-shore or peat bog settlements (Maier 2001). Late Neolithic, Pfyn (Hornstaader Gr.) culture (ca. 5,850–5,450 cal BP). Very rich water-

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

logged remains: einkorn wheat (frequent); emmer wheat (few); naked wheat (prevailing); barley (frequent); pea (few); flax (few); opium poppy (few). Wild: Corylus avellana; Malus sylvestris; Prunus spinosa; Fragaria vesca; Rosa sp.; Quercus sp.; Rubus idaeus/R. fruticosus; Camelina sativa. Aromaric/oil plants: Apium sp.; Anethum graveolens; Petroselinum crispum; Descurainia sophia; Brassica rapa; Galeopsis tetrahit-type. 10. Rosdorf, Lower Saxony (Kirleis and Willerding 2008) Linearbandkeramik culture (6,350±70 uncal BP= ca. 7,450–7,150 cal BP). Einkorn wheat and emmer wheat (co-prevailing); barley (rare); pea (rare); flax (rare); opium poppy (rare). Wild: Corylus avellana.

The Netherlands (General references: Van Zeist 1968–70; Bakels 1991) 1. Graetheide, South Limburg; as well as Sittard, Beek, Elsloo, and Geleen (Bakels 1979, 2001; Bakels and Rousselle 1985). Linearbandkeramik culture (ca. 5,300–5,000 cal BC = ca. 7,250–6,950 cal BP). Numerous remains: einkorn wheat (few); emmer wheat (frequent); flax/linseed (rare); poppy (rare). 2. Maastricht-Randwijck, South Limburg (Bakels et al. 1993). Rössen culture (ca. 4,900–4,600 cal BC = ca. 6,850–6,550 cal BP). Numerous remains (both grains and chaff): einkorn wheat (few); emmer wheat (few); free-threshing wheat (prevailing); naked barely (frequent). Wild: Corylus avellana; Prunus spinosa, and Rosa sp. 3. Swifterbant S3, Flevoland, Western coast, and adjacent sites (van Zeist and Palfenier-Vegter 1981; Cappers and Raemaekers 2008). Early Neolithic (ca. 5,000–3,500 cal BC = ca. 6,950–5,450 cal BP). Numerous remains: einkorn wheat (few); emmer wheat (rare); naked six-rowed and two-rowed barley (prevailing). Wild: Crataegus monogyna; Malus sylvestris; Rubus fruticosus; Corylus avellana; Trapa natans. 4. Aartswoud, North Holland (Pals 1984). Late Neolithic (ca. 2,950–2,750 cal BC = ca. 4,900–4,700 cal BP). Rich remains: einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (few); naked six-rowed barley (prevailing); linseed (few). Wild: Malus sp.; Rubus fruticosus; Corylus avellana; Spergula arvensis; Polygonum convolvulus; P. lapathifolium.

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5. Twisk and Bovenkarspel, West Friesland (Buurman 1987; Buurman et al. 1995). Middle and Late Bronze Age (ca. 1,800–800 cal BC = ca. 3,750– 2,750 cal BP). Numerous remains (both grain and chaff): emmer wheat (frequent); naked barley (prevailing); hulled barley (frequent). Wild: numerous weeds.

Belgium (General references: Bakels 1991) 1. Aubechies, and other contemporary sites (Bakels and Rousselle 1985). Linearbandkeramik culture (ca. 5,400–4,900 cal BC = ca. 7,350–6,850 cal BP). Numerous remains: einkorn wheat (frequent); emmer wheat (prevailing); flax (rare). Wild: numerous herbs. 2. Wange and Overhespen, near St Truiden (Bakels 1992). Linearbandkeramik culture (ca. 5,400–5,000 cal BC = ca. 7,350–6,850 cal BP). Few remains: einkorn wheat (few); emmer wheat (prevailing); naked barley (few); pea (few). Wild: numerous herbs. 3. ‘La Bosse de la Tombe’ à Givry (Heim 1979). Rössen culture (ca. 4,900–4,700 cal BC = ca. 6,850– 6,650 cal BP). Scarce remains: emmer wheat (few); free-threshing wheat (prevailing). Wild: Polygonum hydropiper; Stellaria media; Corylus avellana.

Denmark (General references: Jensen 1991; Robinson 1994, 2003, 2007) 1. Sarup, south-west Funen (Jørgensen 1981). Funnel Beaker culture (ca. 4,350 uncal BP=4,950– 4,850 cal BP). Numerous charred remains: einkorn wheat (rare); emmer wheat (prevailing); free-threshing wheat (rare); naked six-rowed barley (few). Wild: Malus sylvestris; Corylus avellana; Chenopodium album. 2. Norsminde, Krabbesholm, and Visborg, Jutland (Andersen 1991, 2000, 2005; Robinson 2007). Earliest Funnel Beaker culture (ca. 5,850–5,650 cal BP). Few charred remains: emmer wheat (few); naked barley (few). Wild: Pyrus/Malus; Corylus avellana; Glyceria plicata; range of arable weeds, ruderal, and grassland species. 3. Brødrene Gram, Vojens, southern Jutland (Robinson 2000, 2003). (i) Late Neolithic (4,410–

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DOMESTICATION OF PLANTS IN THE OLD WORLD

4,150 uncal BP=ca. 5,050–4,600 cal. BP). Rich charred remains: emmer wheat (frequent); spelt wheat (few); naked barley (prevailing). Wild: arable weed species (few). (ii) Early Bronze Age (ca. 3850–3400 cal BP). Rich charred remains: emmer wheat (few); spelt wheat (few); naked barley (prevailing); hulled six-rowed barley (few). Wild: range of arable weeds, ruderal, and grassland species (abundant). 4. Enkehøj, Gilmosevej, Herning, central Jutland (Møbjerg et al. 2007). Late Neolithic (ca. 3,950–3,650 cal BP). Rich charred remains: emmer wheat (frequent); spelt wheat (few); naked barley (prevailing). Wild: Quercus acorns (abundant); range of arable weeds. 5. Lindebjerg and Voldtofte, Fyn Island (RowleyConwy 1979, 1983). Middle-Late Bronze Age (3,150– 2,850 uncal BP = ca. 3400–2950 cal BP). Rich charred remains: emmer wheat (rare); free-threshing wheat (frequent); hulled barley (frequent); naked barley (prevailing); broomcorn millet (few).

Sweden (General references: Jensen 1991) 1. Settlements in Scania, southern Sweden (Engelmark 1992; Larsson 1992). Early Neolithic (4,925±115 and 4,915±100 uncal BP = ca. 5,900– 5,600 cal BP). Imprints: spelt-type wheats (prevailing); naked barley (frequent); dwarf wheat (few). 2. Settlements in Bjästamon, Ångermanland, northern Sweden (Runesson 2007). Middle Neolithic (3,798 ± 28 uncal BP = ca. 4,650–4,450 cal BP). Hordeum vulgare. 3. Alvastra, Östergötland (Hjelmqvist 1955; Göransson et al. 1995). Middle Neolithic culture (ca. 5,000 cal BP). Numerous carbonized remains and imprints: einkorn wheat (rare); hulled sixrowed barley (rare); naked six-rowed barley (prevailing). Wild: Malus sp. (fruits and pips); Corylus avellana. 4. Eker, Hjulberga, Rosenland, Närke (Hjelmqvist 1979b). Neolithic culture. Scarce charred remains and imprints: emmer wheat (rare); free-threshing wheat (few); naked six-rowed barley (prevailing); pea (rare). Wild: Corylus avellana; Sinapis arvensis.

Norway 1. Hjelle and Gossen, north-west Norway (Soltvedt 2000; Hjelle and Solem 2008). Late Neolithic (Hjelle: 3,760±70 uncal BP = ca. 4,250– 2,000 cal BP; Gossen: 3,72±45 uncal BP = ca. 4,250– 3,950 cal BC). Rich charred remains: naked barley (prevailing, in Hjelle); hulled barley (frequent, in Gossen). 2. Borge vestre, Østfold, south-east Norway (Sandvik 2007, 2008). (i) Late Neolithic (3,680±50 uncal BP = ca. 4,150–3,900 cal BP). Charred grains of barley. (ii) Bronze Age (2,900±30 uncal BP = ca. 3,150–2,950 cal BP). Charred seeds: barley; Panicum miliaceum; flax. 3. Kvåle, south-western Norway (Soltvedt et al. 2007). Late Neolithic (3555±65 uncal BP = ca. 3,900– 3,700 cal BP). Charred remains: emmer wheat (frequent); naked barley (frequent). 4. Forsandmoen, and three adjacent smaller sites (Bakkevig 1982 1995). Bronze Age (ca. 2,900–2,800 cal BP). Rich charred remains: emmer wheat (rare); free-threshing wheat (rare); naked barley (prevailing, and the only identified cereal at two of the sites); hulled barley (rare); oat (rare). Wild: Corylus avellana; Rubus idaeus; Polygonum persicaria; P. tomentosum.

Finland (General references: Jensen 1991; Häkkinen and Lempiäinen 1996) Niuskala, Turku, south-west Finland (Vuorela and Lempiäinen 1988). Late Neolithic Kiukainen culture (3,200±170 uncal BP = 3,600–3,250 cal BP). Numerous remains of charred six-rowed naked barley (recently confirmed by the presence of Hordeum-type pollen grains).

Britain and Ireland (General references: Moffett et al. 1989; Greig 1991; Jones and Rowley-Conwy 2007) 1. Hambledon Hill, Dorset (Jones and Legge 1987, 2008). Early Neolithic (ca. 4,670±80 uncal BP = ca. 5,600–5,050 cal BP). Rich charred remains: emmer wheat, mainly chaff (prevailing); barley, including six-row naked barley (frequent); and a single grape pip. Wild: Corylus avellana.

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

2. Balbridie, Kincardineshire, Grampian, Scotland (Fairweather and Ralston 1993). Neolithic (ca. 4,82±80 uncal BP = ca. 5,700–5,300 cal BP). Numerous charred remains, apparently storage: emmer wheat (prevailing); free-threshing wheat (few, but a pure hoard was found in a posthole); naked barley (frequent); hulled barley (rare); flax (few). 3. The Stumble, Blackwater Estuary, Essex (Murphy 1989). Early Neolithic (ca. 4,675±70 uncal BP = ca. 5,600–5,300 cal BP). Rich, submerged, charred remains: einkorn wheat (few); emmer wheat (prevailing); naked barley (few). Wild: Corylus avellana; Prunus spinosa; Crataegus monogyna; Rubus sp.; Malus sp.; Tilia sp.; Vicia/Lathyrus sp.; Rumex sp.; Polygonum sp.; Galium aparine. 4. Tankardstown, Ireland (Monk 1988). Early Neolithic (4,840±80 uncal BP = ca. 5,650–5,450 cal BP). Numerous charred remains: emmer wheat (prevailing). Wild: Malus sylvestris; Corylus avellana.

France (General references: Marinval 1988; Bakels 1991; Hopf 1991a; Ruas and Marinval 1991; Bouby 2003; Zech-Matterne et al. 2009). 1. La Baume Fontbrégoua, Salernes, Var (Courtin and Erroux 1974). (i) Mesolithic. Few remains: Wild: Vicia sp.; Vitis sylvestris. (ii) Impressed Ware (Cardial) culture (6,180±12 uncal BP; 5,690±130 uncal BP = ca. 7,250–6,250 cal BP). Scarce remains: free-threshing wheat (prevailing). Wild: Lathyrus cf. cicera; Vicia sp.; Lathyrus cf. cicera; Quercus sp. (iii) Middle Neolithic (5680±12 uncal BP; 5,330±12 uncal BP = ca. 6,800–5,800 cal BP). Rich remains: einkorn wheat (few); emmer wheat (prevailing); free-threshing wheat (frequent); six-rowed barley (few); naked six-rowed barley (few). Wild: Vicia sp.; Lathyrus cf. cicera. 2. Pont de Roque Haute, Portiragnes, Hérault (Marinval 2007). Early Neolithic, Impressa culture (6,850±65 uncal BP; 6,745±70 uncal BP= ca. 7,900– 7,500 cal BP). Few carbonized remains: einkorn wheat (few); emmer wheat (few); free-threshing wheat (rare). 3. Châteauneuf-les-Martigues, Font des Pigeons, Bouches-du-Rhône (Courtin et al. 1976). Impressed Ware (Cardial) culture (6,550±100 uncal BP; 6,050±100 uncal BP = ca. 7650–6,700 cal BP). Rich

191

remains: free-threshing wheat and naked six-rowed barley in equal proportions. Wild: Prunus cerasus; Pinus sp. 4. Menneville and several other sites in the Valley of the Aisne (Bakels 1984, 1991; Ilett et al. 1995). Numerous charred remains. (i) Linearbandkeramik culture (6,200±190 uncal BP = ca. 7,550–6,700 cal BP): einkorn wheat (few); emmer wheat (prevailing); naked barley (frequent); pea (rare); lentil (rare). Wild: Corylus avellana; Galium aparine; Bromus sp.; Echinochloa crus-galli; Lapsana communis; Polygonum convolvulus, Rumex sanguinea. (ii) Middle Neolithic, Michelsberg culture: emmer wheat (rare); hulled barley (few); a large number of poppy seeds. 5. Bercy (Paris) (Dietsch 1996, 2001). Middle Neolithic, Chasséen culture (5,130±80 uncal BP; 4,530±12 uncal BP = ca. 6,250–4,900 cal BP). Rich waterlogged and few carbonized remains: emmer wheat (few); free-threshing wheat (frequent); sixrowed barley (few). Wild: Corylus avellana; Vitis sylvestris; Cornus sanguinea; Rubus fruticosus; Quercus sp.; Crataegus monogyna; Rubus idaeus; Prunus spinosa; Sambucus spp.; Malus sylvestris. 6. Grotte G, Baudinard, Var (Courtin and Erroux 1974). Middle Neolithic, Chasséen culture. Numerous remains: emmer wheat (frequent); free-threshing wheat (frequent); six-rowed barley (prevailing); chickpea-like seed (rare); faba bean (rare). Wild: Lathyrus cicera; Vicia sp.; Quercus sp. 7. Station III de Clairvaux, Jura (LundströmBaudais 1984). Late Neolithic (4,412±56 uncal BP; 4,272±70 uncal BP = ca. 5,350–4,650 cal BP) lakeside settlement. Numerous mostly waterlogged remains of threshing and of wild plants: emmer wheat (frequent); free-threshing wheat (few); six-rowed barley (prevailing); flax: seed, capsules, and textile (few); pea (rare); poppy (few). Wild: numerous species including Corylus avellana; Malus sylvestris; Prumus spinosa; Rubus idaeus; R. fruticosus; Fragaria vesca. 8. Grésine, Brison Saint Innocent, Savoie (Bouby and Billaud 2001). Late Bronze Age (Dendrochronology 905–869 BC). Rich waterlogged and carbonized remains: einkorn wheat (few); emmer wheat (frequent); spelt wheat (frequent); free-threshing wheat (few); hulled six-rowed barley (frequent); broomcorn millet (frequent); foxtail millet (prevailing); pea (rare); lentil (rare); faba bean (rare); grass pea

192

DOMESTICATION OF PLANTS IN THE OLD WORLD

(rare); bitter vetch (rare); poppy (frequent); linseed, including capsules (frequent); gold of pleasure, including pods (frequent). Wild: Avena sp.; Brassica campestris; Vitis sylvestris; Quercus sp.; Corylus avellana; Fragaria vesca; Prunus spinosa; Rosa sp.; Rubus spp.; Sambucus spp.

Spain (General references: Hopf 1991a; Zapata et al. 2004; Buxó and Piqué 2008) 1. Balma Margineda, Andorra (Marinval 1995). (i) Mesolithic (9,250±160 uncal BP= ca. 10,650–10,250 cal BP). Wild: Prunus spinosa; Corylus avellana; Quercus sp.; Juglans regia; Pinus pinea. (ii) Early Neolithic (6,850±160 uncal BP = ca. 7,950–5,450 cal BP). Rich remains: emmer wheat (few); free-threshing wheat (frequent); compact type of free-threshing wheat (rare); naked six-rowed barley (few); pea (few). Wild: Quercus sp.; Corylus sp. 2. Coveta de l’Or, Beniarrés, Alicante (Hopf and Schubart 1965; Lopez 1980), and adjacent Cova de Cendres (Buxó, 1997). Early Neolithic Impressed Ware (Cardial) culture (6,340±70 uncal BP = ca. 7,400–7,050 cal BP). Rich remains: einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (frequent); hulled six-rowed barley (frequent); naked six-rowed barley (prevailing); lentil (rare); pea (rare); grass pea (rare); faba bean (rare). Wild: Quercus sp.; Olea europaea var. oleaster. 3. Sites of south-east Spain: (i) Cueva de los Murciélagos, Zuheros, Córdoba (Hopf and Muñoz 1974; Peña-Chocarro 1999). Early Neolithic (6,430±130 to 5,660±12 uncal BP = ca. 7,500–5,660 cal BP). Rich remains: emmer wheat (frequent); freethreshing wheat (prevailing); hulled six-rowed barley (frequent); naked six-rowed barley (frequent); Papaver somniferum seeds. (ii) Cueva del Toro, El Torcal, Antequera, Málaga (Buxó 2004; Hopf unpublished data). Middle Neolithic (5,380±45 uncal BP = ca. 6,300–6,000 cal BP). Rich remains: emmer wheat (few); free-threshing wheat (prevailing); hulled sixrowed barley (few); naked six-rowed barley (frequent); lentil (few); grass pea (few); pea (few); faba bean (frequent); Papaver somniferum seeds. Wild: Quercus acorns; Celtis australis. (iii) Cueva de Nerja, Málaga (Hopf and Pellicer Catalán 1970). Late neolithic (5,065±140 uncal BP = ca. 6,200–5,500 cal BP).

Rich remains: free-threshing wheat (frequent); hulled six-rowed barley (frequent). Wild: Quercus sp.; Olea europaea. 4. Cova de Can Sadurní, Begues, Barcelona (Blasco et al. 1999). (i) Early Neolithic Impressed Ware (Cardial), Late Cardial/Epicardial culture (6405±55 uncal BP to 6,050±110 uncal BP = ca. 7,400– 6,650 cal BP). Rich remains: einkorn wheat (frequent); emmer wheat (frequent); free-threshing wheat (frequent); hulled and naked barley (frequent). Wild: Pistacia lentiscus. (ii) Post-Cardial phases (5,700±110 uncal BP = ca. 6,850–5,800 cal BP). Rich remains: einkorn wheat (few); emmer wheat (frequent); free-threshing wheat (frequent); hulled and naked barley (frequent); pea; lentil; grass pea. Wild: Pistacia lentiscus; Vitis vinifera var. sylvestris; Ficus carica; Quercus sp.; Arbutus unedo; Vicia sativa. 5. North Meseta sites including Cueva de La Vaquera, La Lámpara, and Revilla del Campo (López et al. 2003; Stika 2005; Peña-Chocarro 2007). Early Neolithic (6,12±160 uncal BP = ca. 7,400– 6,650 cal BP). Rich remains: einkorn wheat (rare); emmer wheat (frequent); free-threshing wheat (rare); hulled and naked barley (frequent); Lens sp.; Vicia sativa; Vicia sp. Possibly cultivated: Linum usitatissimum; Papaver somniferum/setigerum. Wild: Vitis vinifera spp. sylvestris; Quercus sp. Weeds from arable fields. 6. Kobaederra, Bizkaia, (Zapata 2002) and El Mirón, Cantabria (Peña-Chocarro et al. 2005). Early Neolithic (5,500±90 uncal BP to 5,375±90 uncal BP = 6,450–5,950 cal BP). Scarce charred remains: einkorn wheat (few); emmer wheat (few); naked wheat (few); barley (few). Wild: Quercus subg. Quercus; Corylus sp.; Rosaceae pomes. 7. Cerro de la Virgen, Orce, Galera, Granada (Buxó 1997). Chalcolitic Period (5,250–4,350 cal BP). (i) Pre-Bell Beaker culture. Few remains: freethreshing wheat (few); hulled barley (few); naked barley (prevailing); faba bean (rare). Wild: rhizome of Stipa tenacissima. (ii) Bell Beaker culture. Rich remains: emmer wheat (few); free-threshing wheat (frequent); compact type of free-threshing wheat (rare); hulled barley (few); naked barley (prevailing); faba bean (few); pea (rare). 8. El Argar culture sites, south-east Spain (Stika 1988; Hopf 1991b). Early Bronze Age (ca. 4,150–3,350

PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES

cal BP). Rich remains: einkorn wheat (few); emmer wheat (few); free-threshing wheat (frequent); hulled six-rowed barley (prevailing); naked six-rowed barley (frequent); pea (few); faba bean (few); flax, seed, and fibers (frequent). Wild: Olea europaea; Vitis vinifera; Ficus carica; Pistacia sp.; Quercus acorns; Celtis australis; Stipa tenacissima.

Portugal (General references: Pinto da Silva 1988; Hopf 1991) 1. Buraco da Pala, Bragança (Ramil Rego and Aira Rodríguez 1993). (i) Late Neolithic (ca. 5,500–4,800 cal BP). Rich remains: free-threshing wheat (frequent); naked six-rowed barley (prevailing); hulled six-rowed barley (rare); faba bean (rare); pea (rare). Wild: Quercus sp. acorns. (ii) Chalcolithic (ca. 4,800– 4200 cal BP). Rich remains: free-threshing wheat (frequent); naked six-rowed barley (prevailing); hulled six-rowed barley (few); pea (rare); lentil (rare); faba

193

bean (few); flax (few); Papaver seeds (few). Wild: Vitis; Quercus sp. acorns; Pinus pinaster, kernel. 2. Zambujal, Torres Vedras (Ramil Rego and Aira Rodríguez 1993). (i) Chalcolithic (ca. 5,400–4,200 cal BP). Scarce remains: einkorn/emmer wheat (rare); free-threshing wheat (few); hulled and naked sixrowed barley (few); faba bean (prevailing); olive (wild? few). Wild: Pinus sp.; Quercus sp. (ii) Bronze Age (ca. 4,000–3,600 cal BP). Numerous remains; einkorn/emmer wheat (rare); free-threshing wheat (frequent); hulled and naked six-rowed barley (frequent); faba bean (prevailing); flax (rare); olive (wild? frequent). Wild: Pinus sp.; Quercus sp. acorns; cork of Quercus suber. 3. Crasto de Palheiros, Murça, Vila Real (Figueiral and Sanches 2003). Chalcolithic (ca. 4,800–4,200 cal BP). Scarce remains, including chaff: emmer/spelt wheat (prevailing); spelt wheat (few); hulled and naked six-rowed barley (few); broomcorn millet (few); faba bean (rare). Wild: Arbutus unedo; Juniperus sp.; Olea europaea.

Appendix A: Site orientation maps

Chokh Imiris-Gora Arukhlo Aratashen and Aknashen

Asikli Höyük

Girikihaciyan

Can Hasan

Çatalhöyük

Jerf el Ahmar Andreas Kastros Kissonerga-Mylouthkia Shillourokambos

Çayönü

Cafer Höyük

Erbaba

Hacilar

Mureybit

Yarym Tepe

Tell Abu Hureyra

Agridhi Khirokitia

Jarmo Tell es-Sawwan

Tell Ramad

Merimde

Tell Aswad Choga Mami

Ohalo Netiv Hagdud-Gilgal Jericho Ghassul 'Ain Ghazal Bab Edh-Dhra

Ali Kosh Tepe Sabz

Saqqara

Fayum

0

100

0

Map 21 South-west Asia, showing the location of the main archaeological sites mentioned in this book.

194

200

200 miles 400 km

Mohelnice BietigheimHeilbronn Bissingen Hilzingen Zürich Egolzwil

Schletz ZánkaVasútállomás

Monte Còvolo

Sammardenchia and Friuli district sites

Sacarovca

Szeged-Gyálarét, Röszke-Lúdvár and Battonya-Basarága Starcevo Liubcova Gomolava Obre

Pienza

Kukoneshti

FüzesabonyGubakút Szilhalom Dévaványa

Mondsee

Sion

Pokrovnik

Poduri

Cîrcea

Sucidava-Delei Karanovo Azmaška Mogila Kapitan Dimitrievo Vršnik Kovacevo Anza Kastanas Nea Nikomedeia

La Marmotta Scamuso

Sesklo

Dimini Grotta dell’Uzzo

Balomenou Lerna Franchthi Cave Knossos

0 0

100 200

200 miles 400 km

Map 22 South and south-east Europe, showing the location of the main archaeological sites mentioned in this book.

Niuskala Borge vestre Rogaland

Balbridie

Alvastra Norsminde, Krabbesholm and Visborg Lindebjerg Sarup

Tankardstown The Stumble Aartswoud

Aubechies

0

100 200

Danau

Swifterbant S3

Hambledon Hill

0

Twisk

Scania region Eker

Graetheide

Kückhoven Rosdorf

Brześć Gniechowice and Stary Zamek

Dresden Ćmielów Menneville and Gniechowice Strachów Aisne Valley LBK sites Nowa Huta-Mogila Bylany G w oździec Mohelnice Bietigheim-Bissingen Hienheim Schletz Hilzingen 200 miles Federseeried Nitriansky Hrádok Blatné Egolzwil 3 Zürich Füzesabony-Gubakút 400 km Wange

Aldenhovener Platte Heilbronn

Map 23 Central and north Europe, showing the location of the main archaeological sites mentioned in this book.

196

APPENDIX A

Aubechies BietigheimMenneville and Aisne Heilbronn Bissingen Valley LBK site Hilzingen Zürich

Egolzwil 3

Sion

Châteauneufles-Martigues El Mirón

Pont de Roque Haute

Kobaederra North Meseta sites Balma Margineda

Buraco da Pala

Cova de Can Sadurní

Cueva de Vaquera La Lámpara& Revilla del Campo Pedra de Ouro Villa Nova de S. Pedro Zambujal

Coveta de l’Or Cova de Cendres

Cueva de los Murciélagos Cerro de la Virgen Cueva del Toro

El Argar culture sites Cueva de Nerja

Kaf Taht El-Ghar

0 0

100

200 miles

200

Map 24 France, Spain, and Portugal, showing the location of the main archaeological sites mentioned in this book.

400 km

Appendix B: Chronological chart for the main geographical regions mentioned in the book

197

BP

South-west Asia

Aegean belt

West Mediterranean

Balkan

Egypt

2,000

BC

0 Iron Age Iron Age

Iron Age

Late Bronze Age

3,000 Late Bronze Age

4,000

5,000

Middle/Late Bronze Age Early/Middle Early/Middle Bronze Age Bronze Age (Minoan, (Otoman) (El Argar) Mycenaean)

Middle Bronze Age

Early Bronze Age

Early Bronze Age

Final Neolithic

New Kingdom

Middle Kingdom

Old Kingdom Chalcolithic

2,000

3,000

Eneolithic (Gumelnitsa, Vancˇa) Archaic

6,000

1,000

Late Neolithic

Middle/Late Neolithic

Middle Neolithic

Early Neolithic Early Neolithic (Impressed (Karanovo, ware) Starcˇevo)

4,000

Chalcolithic

7,000

8,000

Pottery (Ceramic) Neolithic

6,000

Pre-Pottery Neolithic B (PPNB)

10,000

11,000

5,000

Early Neolithic

9,000

Pre-Pottery Neolithic A (PPNA)

Neolithic

e Pr

ing m r - fa

res u t l cu

ic) h t i sol e (M

7,000

8,000

9,000

Fig. 41 Chronological chart for the main geographical regions mentioned in the book. The broken line in each column indicates when definite signs of agriculture start to appear. BP (Before Present) indicates years before 1950, the year 0 of conventional radiocarbon dating.

BP

Central Europe

Ukraine

Alpine Belt

Scandinavia

2,000

BC

0 Iron Age Iron Age

Iron Age

3,000

Bronze Age Bronze Age

Bronze Age

Bronze Age Late Neolithic

4,000

5,000

6,000

Middle/Late Neolithic (Rössen, Lengyel, Funnel Beaker)

Early Neolithic (Linear Bandkeramik)

Late Neolithic

2,000

Early Neolithic (Funnel Beaker)

3,000 Eneolithic (Tripolye) Early Neolithic

Early Neolithic Lake Shore Settlements

4,000

7,000

5,000

8,000

6,000

9,000

10,000

e Pr 11,000

Fig. 41 Continued.

1,000

es ur t l cu g n i rm a f -

) hic t i l eso (M

7,000

8,000

9,000

Appendix C: Information on archaeological sites which appear on Map 2

1: Jeitun (Charles and Hillman 1992; Harris et al. 1993, 1996). 2: Ali Kosh (Helbaek 1969). 3: Jarmo (Helbaek 1959b, 1960, 1966a; Braidwood 1960). 4: Chokh (Lisitsina 1984). 5: Aratashen and Aknashen (Hovsepyan and Willcox 2008). 6: Arukhlo I and II (Januševič 1984; Lisitsina 1984; SchultzeMotel 1988). 7: Çayönü (Van Zeist 1972, 143–66; Van Zeist and De Roller 1991–2, 2003). 8: Aşikli Höyük (van Zeist and de Roller 1995). 9: Tell Abu Hureyra (Hillman 1975, 1989, 2000a; De Moulins 1997, 2000; Hillman et al. 2001). 10: Tell Aswad (van Zeist and Bakker-Heeres 1985). 11: Ain Ghazal (Rollefson et al. 1985). 12: Jericho (Hopf 1983). 13: Yiftah’el (Garfinkel et al. 1988). 14: Kissonerga-Mylouthkia and Shillourokambos (Willcox 2000; Murray 2003). 15: Fayum (Caton-Thompson and Gardner 1934; Wetterstrom 1993; Wendrich and Cappers 2005). 16: Merimde (Wetterstrom 1993). 17: Franchthi Cave (Hansen 1991a, 1992). 18: Nea Nikomdeia (van Zeist and Bottema 1971). 19: Sesklo (Hopf 1962; Kroll 1981a). 2: Knossos (Sarpaki 2009). 21: Rivne (Pashkevich 2003). 22: Sacarovca (Januševič 1984; Kuzminova et al. 1998). 23: Poduri (Cârciumaru and Monah 1985; Monah and Monah 2008). 24: Liubcova (Cârciumaru 1996). 25: Cîrcea (Cârciumaru 1996). 26: Kovacevo (Popova 1992; Marinova 2006). 27: Kapitan Dimitrievo (Marinova 2006, forthcoming). 28: Karanovo (Thanheiser 1997; Marinova 2004, 2006). 29: Azmaška Mogila (Hopf 1973a; Renfrew 1979, Table 7). 30: Anza (Renfrew 1976). 31: Obre (Renfrew 1974). 32: Szeged-Gyálarét, Röszke-Lúdvár and Battonya-Basarága (Hartyányi et al. 1968; Hartyányi and Nováki 1971, 1975; Füzes 1990). 33: Füzesabony-Gubakút (Gyulai 2007). 34: Zánka-Vasútállomás (Füzes 1990, 1991). 35: Bylany (Tempír 1979). 36: Mohelnice (Opravil 1979, 1981; Kühn 1981). 37: Blatné (Hajnalová 1989). 38: Gniechowice and Stary Zamek (Gluza 1994). 39: Strachów (Lityńska-Zając 1997). 40: Brześć (Bieniek 2007). 41: Gwoździec (Bieniek and Litynska-Zajac 2001; LityńskaZając 2007). 42: Pienza (Castelletti 1976). 43: La Marmotta (Rottoli 1993, 2002). 44: Sammardenchia and Friuli district 200

sites (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007). 45: Scamuso (Costantini et al. 1997). 46: Grotta dell’Uzzo (Costantini 1989). 47: Schletz (Kohler-Schneider 2007; Schneider 1994). 48: Egolzwil 3 (Bollinger 1994; Jacomet 2007). 49: Zürich (Jacomet 1988, 2004; Jacomet et al. 1989; Brombacher and Jacomet 1997; Favre 2002; Brombacher et al. 2005). 50: Sion (Martin et al. 2008b; Lundström-Baudais in press). 51: Hilzingen (Stika 1991). 52: Heilbronn (Bertsch and Bertsch 1949; Stika 1996a). 53: Rosdorf (Kirleis and Willerding 2008). 54: Hienheim (Bakels 1978). 55: BietigheimBissingen (Piening 1989). 56: Kückhoven (Knörzer 1998). 57: Aldenhovener Platte (Knörzer 1973, 1974, 1997). 58: Niuskala (Vuorela and Lempiäinen 1988). 59: Scania region (Engelmark 1992; Larsson 1992). 60: Bjästamon (Runesson 2007). 61: Hjelle and Gossen (Soltvedt 2000; Hjelle and Solem 2008). 62: Borge vestre (Sandvik 2007, 2008). 63: Norsminde, Krabbesholm and Visborg (Andersen 1991, 2000, 2005; Robinson 2007). 64: Pont de Roque Haute (Marinval 2007). 65: Graetheide (Bakels 1979, 2001; Bakels and Rousselle 1985). 66: Swifterbant S3 (van Zeist and Palfenier-Vegter 1981; Cappers and Raemaekers 2008). 67: Wange and Overhespen (Bakels 1992). 68: Aubechies (Bakels and Rousselle 1985). 69: The Stumble (Murphy 1989). 70: Hambledon Hil (Jones and Legge 1987, 2008). 71: Balbridie (Fairweather and Ralston 1993). 72: Tankardstown (Monk 1988). 73: Menneville and Aisne Valley LBK sites (Bakels 1984, 1991; Ilett et al. 1995). 74: Châteauneuf-les-Martigues (Courtin et al. 1976). 75: Balma Margineda (Marinval 1995). 76: Coveta de l’Or and Cova de Cendres (Hopf and Schubart 1965; Lopez 1980; Buxó 1997). 77: Cova de Can Sadurní (Blasco et al. 1999). 78: North Meseta sites ( López et al. 2003; Stika 2005; Peña-Chocarro 2007). 79: Cueva de los Murciélagos (Hopf and Muñoz 1974; Peña-Chocarro 1999). 80: Kaf Taht El-Ghar (Ballouche and Marinval 2003). 81: Kobaederra and El Mirón (Zapata 2002; Peña-Chocarro et al. 2005). 82: Buraco da Pala (Ramil Rego and Aira Rodríguez 1993). 83: Zinchecra (van der Veen 1992a, 1992b).

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Index

Page numbers in italic refer to Figures and Tables, and those in bold to Maps Aartswoud 189, 195 Aegilops cylindrica 50 squarrosa 24, 46–49 tauschii 24, 29, 30–33, 46, 46, 47–49, 50 ’Ain Ghazal 2, 194. Colour plate 6 Aknashen 176, 194. Colour plate 6 Albertfalva 183 Aldenhovener Platte 188, 195. Colour plate 6 Ali Kosh 2, 169–70, 194. Colour plate 6 Allium ampeloprasum 156, 174 cepa (onion) 6, 157 longicuspis 157 macrochaetum 157 oschaninii 157 porrum (leek) 6, 7, 155–6 sativum (garlic) 6, 156–7, 156 truncatum 157 tuncelianum 157 vavilovii 157 almond (Amygdalus communis) 6, 116, 147–9. Colour plate 17 archaeological remains 148–9, 149 wild ancestry 147–8 Alvastra 190, 195 amplified fragment length polymorphism (AFLP) 15, 35, 41, 54, 66, 88, Amygdalus bucharica 148 communis see almond fenzliana 148 korschinskyi 147 kuramica 148 persica 144 webbii 148 Anau 176–7 Anethum graveolens (dill) 164 Anza 179, 195. Colour plate 6 Anzingerberg/Hundssteig 183 Apium graveolens (celery) 160–1

apple (Malus domestica) 5, 115, 135–8 archaeological remains 136–8 wild ancestry 135–6. Colour plate 14 distribution map 137 apricot (Armeniaca vulgaris) 7, 116, 144 Aratashen 176, 194. Colour plate 6 archaeological evidence 9–13, 10, 11 availability of 4–5 charred (carbonized) remains 10–11 chemical tests 13 desiccated remains 12 digested remains 12–13 impressions 11–12 mineralization 12 phytoliths 12 preservation by metal oxides 12 south-west Asian Neolithic sites 2 waterlogged preservation 12 Armeniaca vulgaris (apricot) 7, 144 Arukhlo 1 and 2 176, 194. Colour plate 6 Asikli Höyük 2, 171–2, 194. Colour plate 6 asparagus (Asparagus officinalis) 161–2 Atlit Yam 174 Aubechies 189, 195, 196. Colour plate 6 aubergine (Solanum melongena) 8 Austria 183–4 Avena abyssinica 66 byzantina 66, 67 fatua 66, 67 ludoviciana 66, 67 macrocarpa 67 nuda 66, 67 occidentalis 66 sativa see oat sterilis 41, 66, 67, 68. Colour plate 8 strigosa 66

Azmaška Mogila 180, 195. Colour plate 6 Bab edh-Dhra 174, 194 Bajč-Medzi kanálmi 186 Balbridie 191, 195. Colour plate 6 Balma Margineda 192, 196. Colour plate 6 banana (Musa) 8 barley (Hordeum vulgare) 21, 52–8, 54 archaeological remains 1, 56–8, 57 classification 16–17, 57 molecular analysis 51, 55–56 seed dispersal 55 six-rowed cultivated barley 55 two-rowed cultivated barley 52–3 wild progenitor 3, 4, 53–6, 55 distribution 55–6, 57 Battonya-Basarága 182, 195. Colour plate 6 beet (Beta vulgaris) 7, 159–60 Belgium 189 Bercy 191 Berettyóújfalu-Szilhalom 182, 195 Beta adanensis 159 macrocarpa 159 maritima 159 vulgaris (beet) 7, 159–60 Bietigheim-Bissingen 188, 195, 196. Colour plate 6 bitter orange (Citrus aurantium) 7, 146–7 bitter vetch (Vicia envilia) 1–2, 2, 61, 75–76, 79, 92–5 archaeological remains 76, 92–5, 94 wild progenitor 4, 92 distribution 93 Bizkaia 192 Bjästamon 190. Colour plate 6 black cumin (Nigella sativa) 164–5 Blatné 186, 195. Colour plate 6 Bölcske-Vörösgyír 183 Borge Vestre 190, 195. Colour plate 6 237

238

INDEX

bottle gourd (Lagenaria siceraria) 155 Bovenkarspel 189 Brassica bourgeaui 158 campestris 112, 159 cretica 158 hilarionis 158 insularis 158 juncea 112 macrocarpa 67, 158–9 napus (swede) 112, 159 nigra 112, 181 oleracea (cabbage) 158–9 rapa (turnip/oil-seed rape) 7, 112, 159 rupestris 158 villosa 158 bread wheat (Triticum aestivum) 24, 29, 47–51 archaeological remains 50–1 club wheat 28, 29, 48 cytogenetics 47 free-threshing bread wheats 31, 33, 48, 51 hulled bread wheats 30, 31–2, 48, 50–1 molecular analysis 50 spelt wheat 29, 31–2, 48, 50–1 wild ancestry 48–50 see also wheats Britain 190–1 broad bean see faba bean broccoli 158 Brødrene Gram 189–90 broomcorn (common) millet (Panicum miliaceum) 5, 7, 20, 69–71. Colour plate 9 archaeological remains 69–71, 70 Brussels sprout (Brassica oleracea) 158 Brześć 185, 195. Colour plate 6 Budapest 183 Bulgaria 179–81 Buraco da Pala 193, 196. Colour plate 6 Bylany 186, 195. Colour plate 6 cabbage (Brassica oleracea) 158–9 Cafer Höyük 2, 194 Camelina microcarpa 111 sativa see gold of pleasure Can Hasan III 171, 194 Cannabis indica 106 sativus see hemp Canton of Grisons 187 Canton of Valais 187 Cap Andreas-Kastros 177, 194

caprification 128 carob (Ceratonia siliqua) 145–6 archaeological remains 145–6 reproductive biology 145 carrot (Daucus carota) 7, 160 Carthamus flavescens 168 gypsicola 168 oxyacanthus 168 palaestinus 168 persicus 168 tinctorius (safflower) 168 Castanea sativa (chestnut) 150–1 Çatalhöyük 171, 194 Caucasia 176 Çayönü 2, 171, 194. Colour plate 6 Celei (Sucidava) 181 celery (Apium graveolens) 160–1 Ceratonia siliqua see carob cereals 21–74 see also specific plants Cerro de la Virgen 192, 196 Chateauneuf-les-Martigues 191, 196. Colour plate 6 cherries (Prunus avium and P. cerasus) 143–4 archaeological remains 144 sour cherry 116, 143 sweet cherry 115, 143 chestnut (Castanea sativa) 150–1 chickpea (Cicer arietinum) 75, 87–9 archaeological remains 1–2, 2, 76, 88–9 molecular analysis 88 seed dispersal 87 wild ancestry 3–4, 87–8 distribution map 88 Choga Mami 171, 194 Chokh 176, 194. Colour plate 6 chronological chart 197–9 chufa (Cyperus esculentus) 5, 6, 158 Cicer arietinum see chick pea bijugum 87 echinospermum 87 judaicum 87, 89 pinnatifidum 87 reticulatum 87, 88, 89. Colour plate 10 Circea (Cârcea) 181, 195. Colour plate 6 citron (Citrus medica) 7, 146 Citrullus colocynthis 154 ecirrhosus 154 lanatus see watermelon rehmii 154 vulgaris 153

Citrus aurantiifolia 7, 147 aurantium (bitter orange) 7, 146–7 limon (lemon) 7, 146 maxima (pumello) 7, 146 medica 7, 146 paradisi 147 reproductive biology 146 reticulata 147 citrus fruits 146–7 Ćmielów 185, 195 Colocasia esculenta (taro) 8 common millet see broomcorn millet common oat see oat common vetch (Vicia sativa) 95 archaeological remains 95 condiments 163–5 see also specific plants coprolites 12–13, 108 coriander (Coriandrum sativum) 163–4 Corylus avellana (hazelnut) 151 maxima 151 cottons (Gossypium arboreum and G. herbaceum) 7, 100, 107–9 archaeological remains 108–9 genetic analysis 108 Cova de Can Sadurní 192, 196. Colour plate 6 Cova de Cendres 192, 196. Colour plate 6 Coveta de l’Or 192, 196. Colour plate 6 cow pea (Vigna unguiculata) 8, 73 crab apples (Malus sylvestris) 135–7 distribution 137 Crasto de Palheiros 193 Crete 179 Crocus cartwrightianus 165 sativus (saffron) 165 Csepel-Vízmü 183 Cuconeştii Vechi 182 cucumber (Cucumis sativus) 7, 155 Cucumis callosus 155 chate 155 melo see melon sativus (cucumber) 7, 155 Cueva de La Vaquera 192, 196 Cueva de los Murciélagos 192, 196. Colour plate 6 Cueva del Toro 192, 196 Cueva de Nerja 192, 196 cumin (Cuminum cyminum) 163, 164 black cumin 163, 164–5

INDEX

Cydonia oblonga (quince) 144–5 vulgaris 144–5 Cyperus esculentus (chufa) 5, 158 Cyprus 177 cytogenetics 14 see also specific plants Czech Republic 185–6 damson 140–141 Dannau 188, 195 date palm (Phoenix dactylifera) 5, 114, 116, 131–4 archaeological remains 115, 133–4, 133 molecular analysis 133 reproductive biology 131 wild progenitor 131–3 distribution map 132 Daucus carota (carrot) 7, 160 dendrochronology 17–18 Denmark 189–90 Dévaványa Réhelyi 182, 195 Dhali Agridhi 177 dill (Anethum graveolens) 163, 164 Dimini 178, 195 disarticulation scars 1, 22, 26, 27, 31–2, 53, 54, 116, Djade el Mughara 173 DNA analysis 13–15, 32, 35, domestication syndrome 9, 13 cereals 23 pulses 75–7 Dresden-Nickern 188, 195 Dürrnberg/Hallein 183–4 durum wheat see emmer and durumtype wheats (Triticum turgidum) dye crops 166–8 see also specific plants dyer’s rocket (Reseda luteola) 167 Eberdingen-Hochdorf 188 Egolzwil 3 186, 195, 196. Colour plate 6 Egypt 174–6 einkorn wheat (Triticum monococcum) 24, 25, 29, 34–40 archaeological remains 1, 32, 36–40, 37, 38 beginnings of domestication 1 grain dispersal 31, 35 molecular analysis 35 wild progenitor 4, 26, 34–6, 33 distribution 35, 36 see also wheats Eker 190, 195 El Argar 192–3, 196 El Mirón 192, 196. Colour plate 6

emmer and durum-type wheats (Triticum turgidum) 23, 24, 26, 27, 29, 40–7. Colour plate 4 archaeological remains 1, 42–7, 43, 45 hulled emmer wheats 41–47, 43, 45. Colour plate 5 free-threshing wheats 46–7 characterization of cultivated forms 26, 27, 29, 40, cytogenetics 41 grain dispersal 31, 41. Colour plate 2, 3 molecular analysis 41, 42 wild progenitor 3, 26, 27, 34, 40–2. Colour plate 1, 2 distribution 16, 41–2, 42 see also wheats Enkehøj 190 enzyme polymorphism 15 Erbaba 172, 194 faba (broad) bean (Vicia faba) 75, 89–92. Colour plate 11 archaeological remains 76, 91–2, 91 characterization of cultivated forms 90 cytogenetics 90 molecular analysis 90 wild ancestry 90–1 Farafra Oasis 174 Fayum 175, 194. Colour plate 6 Federseeried 188, 195 fenugreek (Trigonella foenum-graecum) 75, 97–8 archaeological remains 98 fibre-producing crops 100–13 see also specific plants Ficus carica see fig colchica 128 geraniifolia 128 hyrcanica 128 johannis 128 palmata 128 pseudosycomorus 128 sycomorus see sycamore fig virgata 128 fig (Ficus carica) 5, 6, 114, 116, 126–30, 127 archaeological remains 115, 129–30 reproductive biology 126–8 wild ancestry 128–9 distribution map 129 Finland 190 flax (Linum usitatissimum) 1, 2, 61, 100, 101–6. Colour plate 12 archaeological remains 2, 103–6, 104, 105

239

molecular analysis 101–3 wild ancestry 3, 101–3, 102 distribution map 103 Forsandmoen 190 foxtail (Italian) millet (Setaria italica) 5, 7, 71–2. Colour plate 9 archaeological remains 70, 71–2 seed dispersal 71 wild progenitor 71 France 191–2 Franchthi Cave 177–8, 195. Colour plate 6 fruit crops 5–6, 114–16 see also specific species Füzesabony-Gubakút 182, 195. Colour plate 6 Galabovo 180–1 garlic (Allium sativum) 6, 156–7, 156 Germany 188–9 germination inhibition 22, 76, 82, 100 Ghassul see Tuleilat Ghassul Gilgal 173 Gilmosevej 190 Girikihaciyan 172, 194 gluten proteins 23 Gniechowice 185, 195. Colour plate 6 gold of pleasure (Camelina sativa) 7, 16, 100, 111 Gomolava 179, 195 Gossen 190 Gossypium arboreum see cottons barbadense 107 herbaceum see cottons hirsutum 107 Graetheide 189, 195. Colour plate 6 grafting 5, 6, 114–115 see also specific plants grain size 22 grapefruit (Citrus paradisi) 146, 147 grapevine (Vitis vinifera) 5, 6, 114, 116, 121–6 archaeological remains 124–6 pip morphology 122–3, 123, 124–5 reproductive biology 124 sex determination 124 wild progenitor 121–4, 122 distribution 122, 123 grass pea (Lathyrus sutivus) 2, 95–6 archaeological remains 96, 97 cytogenetics 95 Greece 177–9 greengage 141 Grésine 191–2 Grotta dell’Uzzo 184, 195. Colour plate 6

240

INDEX

Grotte G, Baudinard 191 Gwoździec 185. Colour plate 6 Hacilar 171, 194 Hala Sultan Tekke 177 Hambledon Hill 190, 195. Colour plate 6 hazelnut (Corylus avellana) 151 Heilbronn 188, 195, 196. Colour plate 6 hemp (Cannabis sativa) 7, 100, 106–7 Hienheim 188, 195. Colour plate 6 Hierakonpolis 175 Hilzingen 188, 195, 196. Colour plate 6 Hjelle 190. Colour plate 6 Hordeum agriocrithon 53, 57 distichum 52, 53, 55, 57 hexastichum 52, 57 spontaneum 41, 53–9, 54, 55, 56. Colour Plate 157 tetrastichum 52 vulgare see barley Hornstaad 188–9 horse bean see faba bean horticulture 5–6, 114–17 see also specific plants Hungary 182–3 hyacinth bean (Lablab purpureus) 73 Imiris-Gora 176, 194 Indigofera 166–168 arrecta 168 tinctoria (indigo) 7, 166, 168 Indigo (Indigofera tinctoria) 7, 166–168 Iran 169–70 Iraq 170–1 Ireland 190–1 Isatis tinctoria (woad) 166–7 Israel 173–4 Italian millet see foxtail millet Italy 184–5 Jarmo 2, 170, 194. Colour plate 6 Jeitun 176. Colour plate 6 Jerf el Ahmar 173, 194 Jericho 2, 174, 194. Colour plate 6 Jerma 176 Jordan 173–4 Juglans regia (walnut) 149–50 Kaf Taht el-Ghar 176, 196. Colour plate 6 kale 158–9 Kapitan Dimitrievo 180, 195. Colour plate 6 Karanovo 179–80, 195. Colour plate 6

Kastanas 178–9, 195 Khirokitia 177, 194 Kissonerga-Mylouthkia 2, 177, 194. Colour plate 6 Knossos 179, 195. Colour plate 6 Kobaederra 192, 196. Colour plate 6 kohlrabi 158 Kom el-Hisn 175 Kovacevo 180, 195 Krabbesholm 189, 195. Colour plate 6 Kückhoven 188, 195. Colour plate 6 Kukoneshti 195 Kumtepe 172 Kvåle 190 La Baume Fontbrégoua 191 Lablab purpureus (hyacinth bean) 73 ’La Bosse de la Tombe’ à Givry 189 Lactuca sativa (lettuce) 6, 7, 157–8 serriola 157 Lagenaria siceraria (bottle gourd) 155 Lake Biel 187 Lake Neuchatel 187 La Lámpara 192, 196 La Marmotta 185, 195. Colour plate 6 Lathyrus cicera 95–6 clymenum (Spanish vechling) 97 sativus see grass pea leek (Allium portum) 6, 7, 155–6 legumes 75–7 see also specific plants lemon balm (Melissa officinalis) 163 lemon (Citrus limon) 7, 146–7 Lens culinaris see lentil ervoides 77 lamottei 77 nigricans 77 odemensis 78 orientalis 77–9, 78, 80 tomentosus 77, 78 lentil (Lens culinaris) 75, 77–82 archaeological remains 1–2, 76, 79–82, 80 cytogenetics 77–9 identification of cultivated forms 81 molecular analysis 78 seed dispersal 79 wild progenitor 4, 77–9, 79 distribution 78 Lerna 178, 195 lettuce (Lactuca sativa) 6, 7, 153, 157–8 Libya 176 lime (Citrus aurantiifolia) 7, 146–7 Lindebjerg 190, 195 linseed see flax

Linum angustifolium 101 bienne 101, 102 usitatissimum see flax Liubcova 181, 195. Colour plate 6 Luca Vrublevecaja 182 lupins (Lupinus) 98–9 albus (white lupin) 98 wild progenitor 98–9, 98 angustifolius 98 graecus 99 luteus 98 termis 98 Maadi 174–5 Maastricht-Randwijck 189 madder (Rubia tinctorum) 166, 167–8 Majaki 182 Majdaneckoe 182 Malus domestica see apple kirghizorum 136 orientalis 136. Colour plate 14a prunifolia 136 pumila see apple sieversii 136. Colour plate 14b sylvestris 136. Colour plate 14a turkmenorum 136 mandarin (Citrus reticulata) 146, 147 marrowstem kale 158 Melissa officinalis (lemon balm) 163 melon (Cucumis melo) 6, 154–5 archaeological remains 155 Menneville and Aisne Valley sites 191, 195, 196. Colour plate 6 Merimde 175, 194. Colour plate 6 microsatellites (SSR) 15, 84, 119 migrants temperate crops from central and/ or east Asia 7 warm-weather crops from south and/or east Asia 7–8 warm-weather crops from sub-Saharan Africa 8 millets see broomcorn millet; foxtail millet mineralization 12 Mohelnice 185–6, 195. Colour plate 6 Moldavia 181–2 molecular biology tools 14–15 recessive mutations 61 see also specific plants Mondsee 183, 195 Monte Còvolo 184–5, 195 Morocco 176 Musa (banana and plantain) 8 mustard 112

INDEX

Nabta Playa 174 Nahal Zehora II 173 Naqada 175 Nea Nikomedeia 178, 195. Colour plate 6 Neolithic south-west Asian crop assemblage 1–3, 3 spread 3, 4. Colour plate 6 Netherlands 189 Netiv Hagdud 173, 194 Nigella sativa (black cumin) 164–5 Nitriansky Hrádok 186, 195 nitrogen fixation 75, 97 Niuskala 190, 195. Colour plate 6 Norsminde 189, 195. Colour plate 6 Norway 190 Novye Rusešty 182 Nowa Huta-Mogila 185, 195 oat (Avena sativa) 7, 16, 21, 66–9 archaeological remains 69, 70 cytogenetics 66 molecular analysis 66 seed dispersal 67–8 wild ancestry 66–7, 68 Obre 179, 195. Colour plate 6 Ohalo II 173, 194 oil-producing crops 100 oil-seed rape (Brassica rapa) 112, 159 Olea europaea see olive oleaster 116, 117–18. Colour plate 13 olive (Olea europaea) 5, 6, 114, 116–21. Colour plate 13 archaeological remains 115, 118, 119–21 grafting 117–18 molecular analysis 119 vegetative propagation 117 wild progenitor 117–19. Colour plate 13 distribution 117–19, 119 onion (Allium cepa) 6, 157 orange 146, 147 Seville or bitter orange (Citrus aurantium) 7, 146–7 Oryza nivara 74 rufipogon 74 sativa see rice Overhespen 189. Colour plate 6 Panicum milliaceum see broomcorn millet spontaneum 69 Papaver setigerum 109, 110, 111 somniferum see poppy

parsley 163 parsnip (Pastinaca sativa) 153, 161 parthenocarpy 116, 127, 130 peach (Persica vulgaris) 7, 116, 144–5 pea (Pisum sativum) 75, 82–7, 84 archaeological remains 1, 2, 76, 84–7, 85 cytogenetics 82, 83–4 molecular analysis 84 wild ancestry 4, 82–4 distribution map 83 morphological types 82–3 pearl millet (Pennisetum glaucum) 8, 72, 73 pear (Pyrus communis) 5, 115, 138–40 archaeological remains 140 wild ancestry 138–40 distribution map 139 Pennisetum glaucum see pearl millet Persica vulgaris (peach) 7, 116, 144–5 Phoenix dactylifera see date palm reclinata 133 sylvestris 133 theophrastii 133 Pienza 184, 195. Colour plate 6 pistachio (Pistacia vera) 7, 115, 116, 151–2 Pistacia atlantica 151 khinjuk 151 palaestina 151 terebinthus 151 vera (pistachio) 7, 115, 116, 151–2 Pisum elatius 82–3, 84 fulvum 82 humile 83, 84 sativum see pea syriacum 83 plantain (Musa) 8 plum (Prunus domesticus) 5, 115, 116, 140–3 archaeological remains 142–3, 143 cytogenetics 141, 142 wild ancestry 141–2 Poduri 181, 195. Colour plate 6 Pokrovnik 179, 195 Poland 185 Polish wheat 29, 39 polymerase chain reaction (PCR) 15 polyploidy 14 see also specific plants pomegranate (Punica granatum) 6, 115, 134–5 archaeological remains 134–5

241

Pont de Roque Haute 191, 196. Colour plate 6 poppy (Papaver somniferum) 5, 109–11, 110 archaeological remains 109–11 wild poppy distribution map 110 Portugal 193 Pre-Pottery Neolithic B (PPNB) 1, 2–3, 42, 43 preservation of plant remains 9–13, 11 carbonization 10–11 desiccation 12 digested remains 12–13 impressions 11–12 metal oxides 12 mineralization 12 waterlogging 12 protein variation 15 Prunus amygdalus 147 armeniaca 144 avium see cherries brigantina 142 caspica 142 cerasifera 141–2 cerasus see cherries cocomilia 142 divaricata 141 domestica see plum dulcis 147 fruticosa 143 insititia 141 persica 144 salicina 141 sogdiana 142 spinosa 141, 142 ursina 142 pulses 75–7 see also specific plants pummelo (Citrus maxima) 7, 146, 147 Punica granatum see pomegranate Putineşti 182 Pyrus amygdaliformis 140 bretschneideri 140 caucasica 139, 140 communis see pear domestica 138 elaeagrifolia 140 korshinskyi 140 pyraster 139, 140 pyrifolia 140 salicifolia 140 spinosa 139–40 syriaca 140. Colour plate 15, 16 ussuriensis 140 quince (Cydonia vulgaris) 144, 145

242

INDEX

radiocarbon dating 17–19, 18, 19 radish (Raphanus sativus) 112 Rákoskeresztúr-Újmajor 182 random amplified polymorphic DNA (RAPD) 15, 66, 101, 119 recessive mutations 60 Reseda luteola (dyer’s rocket) 167 restriction fragment length polymorphism (RFLP) 15, 41 Revilla del Campo 192, 196 rice (Oryza sativa) 5, 7, 21, 73–4 Rivne 182. Colour plate 6 Rosdorf 189, 195. Colour plate 6 Röszke-Lúdvár 182, 195. Colour plate 6 Rovno see Rivne Rubia tinctorum (madder) 167–8 Rumania 181 Ruseştii Noi 182 rush nut see chufa rye (Secale cereale) 7, 59–65, 63 archaeological remains 2, 37, 64–5, 64 molecular analysis 59 wild ancestry 59–62 distribution map 62 Sacarovca 181–2, 195. Colour plate 6 Saccharum officinarum (sugar cane) 7–8 safflower (Carthamus tinctorius) 166, 168 saffron (Crocus sativus) 165 Sammardenchia 184, 195. Colour plate 6 San Marco 184 Saqqara 175, 194 Šarišské Michalˇany-Fedelemka 186 Sarup 189, 195 Scamuso 184, 195. Colour plate 6 Scania 2, 190, 195. Colour plate 6 Schletz 183, 195. Colour plate 6 Secale afghanicum 59 anatolicum 61 ancestrale 59, 60, 63 cereale see rye ciliatoglume 61 dalmaticum 61 dighoricum 59 iranicum 60, 61 kupriyanowii 61 montanum 61, 63–4, 66. Colour plate 7 segetale 59 strictum 61 sylvestre 60, 62

vavilovii 59, 61, 62 secondary crops 7 seed dispersal cereals 22–3 barley 53 foxtail millet 71 oats 66–8 rye 60 wheats 30–1, 35, 40 pulses 76 chickpea 87 lentil 79 flax 106 self-pollination 20–1 sesame (Sesamum indicum) 7, 112–13 archaeological records 113 Sesamum indicum see sesame orientale 112 Sesklo 178, 195. Colour plate 6 Setaria italica see foxtail millet viridis 71 Seville or bitter orange (Citrus aurantium) 7, 146–7 Shillourokambos 2, 177, 194 Shiqmim 174 Shortugai 177 Sion 187, 195. Colour plate 6 simple sequence repeat (SSR) 15, 84, 119 Sinapis alba (white mustard) 112 single nucleotide polymorphisms (SNPs) 15 sloe (Prunus spinosa) 141, 142 Slovakia 185–6 Solanum melongena (aubergine) 8 sorghum (Sorghum bicolor) 8, 20, 72–3 south-west Asian crop assemblage 1–4 spread of 3, 4. Colour plate 6 Spain 192–3 Spanish vechling (Lathyrus clymenum) 97 spelt wheat, see bread wheat Starčevo 179, 195 Starye Kukoneshti 182 Stary Zamek 185, 195 Station III de Clairvaux 191 Stillfried 183 Strachów 185, 195. Colour plate 6 Sucidava-Celei 181, 195 sugar cane (Saccharum officinarum) 7 swede (Brassica napus) 112, 159 Sweden 190 Swifterbant S3 189, 195. Colour plate 6 Switzerland 186–7 sycamore fig (Ficus sycomorus) 115, 116, 130–1

archaeological remains 130–1 reproductive biology 130 Syria 172–3 Szeged-Gyálarét 182, 195. Colour plate 6 Szilhalom 195 Tankardstown 191, 195. Colour plate 6 taro (Colocasia esculenta) 8 Tell Abu Hureyra 2, 172, 194. Colour plate 6 Tell Aswad 2, 173, 194. Colour plate 6 Tell es-Sa’idiyeh 174 Tell es-Sawwan 170, 194 Tell Hârşova 181 Tell Mureybit 172–3, 194 Tell Ramad 194 Tepe Hasanlu 170 Tepe Sabz 170, 194 Tepe Yahya 170 The Stumble 191, 195. Colour plate 6 Timopheev’s wheat (Triticum timopheevi) 24, 29, 42, 51 cultivated 24, 51 cytogenetics 51 wild 41, 51 see also wheats Toos-Waldi 187 Toumba Balomenou 178, 195 Transcaucasia 176 Trigonella foenum-graecum see fenugreek Triticum aegilopoides 39, 33, 34, 35 aestivum see bread wheat araraticum 29, 42, 51 baeoticum see einkorn wheat carthlicum 29, 40 compactum 29, 48 dicoccoides see emmer and durum-type wheats dicoccum see emmer and durum-type wheats durum see emmer and durum-type wheats macha 29, 47 monococcum see einkorn wheat parvicoccum 29, 40 polonicum 29, 40 sativum 29, 48 sinskajae 29, 34 spelta see bread wheat sphaerococcum 29, 48 thaoudar 29, 35 timopheevi see timopheev’s wheat turgidum see emmer and durum-type wheats

INDEX

urartu 23–4, 29, 36, 41 vulgare 29, 48 Troy 172 Tuleilat Ghassul 174, 194 Turkey 171–2 turnip (Brassica rapa) 7, 112, 153, 159 Tutankhamun tomb 175–6 Twisk 189, 195 Ukraine 181–2 Usatovo 182

narbonensis 90, 91, 92 sativa see common vetch serratifolia 90 Vigna unguiculata see cow pea Visborg 189, 195. Colour plate 6 viticulture see grapevine Vitis sylvestris 121, 122, 123 vinifera see grape vine Voldtofte 190 Vršnik 179, 195

vegetables 6–7, 153 see also specific plants vegetative propagation 114–15 carob 145 fig 126–7 grapevine 124 olive 117 vetch see bitter vetch; common vetch Vicia ervilia see bitter vetch faba see faba bean galilea 90, 91 hyaenis-ciamus 90 johannis 90 kalakhensis 90

walnut (Juglans regia) 149–50 Wange 189, 195. Colour plate 6 watermelon (Citrullus lanatus) 6, 153–4 archaeological remains 154 weeds 7, 10, 16 oil plants 100 wheats (Triticum) 20, 23–32 beginnings of domestication 1 classification 16, 23–4, 29 cytogenetics 23–4 free-threshing wheats 24–32, 27, 28, 31, 32 tetraploid vs hexaploid free-threshing wheats 32, 32

243

hulled wheats 24–30, 26, 30, 31 wild progenitors 3–4, 13–16, 33 see also bread wheat; einkorn wheat; emmer and durum-type wheats; Timopheev’s wheat white lupin (Lupinus albus) 98 wild progenitor 98–9, 98 wild progenitors 3–4 geographic distribution 3–4, 15–16 identification 13–15 classical taxonomic methods 14 cytogenetic analysis 14 molecular analysis 14–15 see also specific plants woad (Isatis tinctoria) 166–7 Yarym Tepe 170–1, 194 Yiftah’el 2. Colour plate 6 Yugoslavia (former) 179 Yunatzite 181 Zambujal 193, 196 Zánka-Vasútállomás 182, 195. Colour plate 6 Zinchecra 176. Colour plate 6 Zürich 186–7, 195, 196. Colour plate 6

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Plate 1 Stands of wild emmer wheat, Triticum dicoccoides (upper), and wild barley, Hordeum spontaneum (lower)—the progenitors of domesticated emmer wheat and domesticated barley (photographs kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).

Plate 2 A spikelet of wild tetraploid emmer wheat, T. turgidum subsp dicoccoides. Note the smooth disarticulation scars, both below and above the internode. These smooth scars are the diagnostic elements for the identification of the wild forms.

Plate 3 A spikelet of domesticated, non-shattering, tetraploid emmer wheat, T. turgidum subsp. dicoccum. Note the rough disarticulation scars, both below and above the internode. These rough scars are the most reliable diagnostic elements for the identification of the domesticated forms.

Plate 4 An ear of domesticated, non-shattering, tetraploid emmer wheat, T. turgidum subsp. dicoccum.

Plate 5 A–Remains of emmer wheat ears retrieved from Saqqara, Lower Egypt. B–Details of two individual ears (courtesy of Agropolis-Museum, Montpellier, France).

60

Legened

61

einkorn wheat emmer wheat barley flax lentil pea

58 62

71

2,500-2,000 BP 3,000-2,500 BP 3,500-3,000 BP 4,000-3,500 BP 4,500-4,000 BP 5,000-4,500 BP 5,500-5,000 BP 6,000-5,500 BP 6,500-6,000 BP 7,000-6,500 BP 7,500-7,000 BP 8,000-7,500 BP 8,500-8,000 BP 9,000-8,500 BP 9,500-9,000 BP 10,000-9,500 BP 10,500-10,000 BP

63 59

72

70

66

69

65 68

40

56

67

53

57

39

52,55

73

37 36

54 47

49

50 82

21

35

51 48

38

44 75

64

22

33

34

81 78

41

1

23

32

4

74

25 6 31

77

24

29

42 43

79

7

27 45

76

5

28 30

26

18 8

19

80

3

46

9

17

2

14

20

10 13 11 12 16

0 0

Scale 1:16,000,000 125 250 375 250

15

500 miles

500 km 83

Plate 6 (Map 2) The spread of the south-west Asian Neolithic crop assemblage in Europe, west Asia, and North Africa. For details on the numbered sites, see Appendix C (p. 200). These are the earliest sites in which domesticated grain crops were found, in each country.

Plate 7 Unripe ear of wild perennial rye, Secale montanum (photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).

Plate 8 A seed dispersal unit of wild oat, Avena sativa subsp. sterilis. Note the smooth disarticulation scar in the lower tip of the spikelet, and the drill-like awans, which helps in the insertion of the spikelet into the ground.

Plate 9 Millet crops of southwest Asia. A and B–Broomcorn millet [= common millet], Panicum miliaceum; A–Fruiting panicle; B–Ripe spikelets. C and D–Foxtail millet [= Italian millet], Setaria italica; C–Fruiting panicle; D–Ripe spikelets (after Vaughan and Geissler 2009; with kind permission of Oxford University Press).

Plate 10 Fruiting branch of wild chickpea, Cicer arietinum subsp. reticulatum.

Plate 11 Faba bean, Vicia faba. A–plant, B–flowers, C–opened pod, D–seed (after Vaughan and Geissler 2009; with kind permission of Oxford University Press).

Plate 12 Flax stems, heckled flax line, and spun linen thread, the work of Alverna Messersmith Learn, early 1900s (with kind permission of Linda Learn, Class Act Fabrics).

Plate 13 Fruiting branches and stones of wild olive, Olea europaea subsp. oleaster (left), and domesticated olive, Olea europaea subsp. europaea (right).

Plate 14A Fruiting branches of wild apple, Malus sylvestris subsp. orientalis (photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).

Plate 14B Flowering branches of wild apple, Malus sieversii (photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).

Plate 15 Flowering branch of wild pear, Pyrus syriaca.

Plate 16 Wild pear, Pyrus syriaca: cluster of maturing fruits (photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).

Plate 17 Almond, Amygdalus communis: cluster of flowers (photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).